Effect of plant species on shrub fertile island at an oasis–desert ecotone in the South Junggar Basin, China

Shrub fertile islands are a common feature in arid ecosystems. To examine the effect of plant species on the spatial patterns of soil chemical and physical properties surrounding individual shrubs, two deciduous shrub species with different morphologies (Tamarix spp. and Haloxylon ammodendron Bge.) were studied at an oasis–desert ecotone in South Junggar Basin. Soil samples were collected under the shrub crown (canopy), at the vertically projected limit of shrub crown margin (periphery), and in the space between shrub crowns (interspace) at two depths, 0–10 and 10–20 cm, to analyze their physical and chemical properties. The results show that the fertile islands of Tamarix spp. are enriched with more soil nutrients (significantly higher, P<0.05; soil organic matter (SOM); total nitrogen (TN) and available nitrogen (AN); to a deeper depth (>20 cm) and in a larger area (beyond the canopied area) compared to that of H. ammodendron (significantly higher, P<0.05, soil nutrients detected only for AN; <20 cm in depth; smaller than the canopied area). Soil texture patterns surrounding the shrubs of the two species are even more different, with more coarse particles under the Tamarix spp. canopies compared to the interspace between shrubs but fewer under the H. ammodendron canopies compared to the interspaces. These variations are attributed to the difference in morphology of the two studied species: the Y-shaped crowns of H. ammodendron are less capable of capturing and maintaining litter under them than the hemispheroidal crowns of Tamarix spp., which leads to the less well developed fertile islands surrounding H. ammodendron shrubs.     

Contrasting nutrient-capture strategies in shrubs and grasses of a Patagonian arid ecosystem

Shallow-rooted grasses and deep-rooted shrubs dominate arid ecosystems where nitrogen is concentrated in the upper layers of the soil and water is distributed throughout. Analysis of mineral nitrogen and absorption patterns using a tracer indicated that shrubs in Patagonia absorbed nutrients from the lower, relatively nutrient-poor layers of the soil. Are they, consequently, at a competitive disadvantage with grasses that have the opposite pattern? Studies of nitrogen economy indicated that shrub and grass species have similar N-use efficiency but that they achieve it through opposite mechanisms. Shrubs have a conservative N economy absorbing annually only small fraction of their N content, whereas grasses have a more open N economy. This study about N-capture strategies in conjunction with previous studies about water-use by shrubs and grasses in the Patagonian Steppe suggest a coupling of N and water-capture strategies. Our findings have implications for the response of arid and semiarid ecosystems to global warming, nitrogen deposition, and biodiversity change. For example, climate change scenarios predict, for most arid regions, decreases in moisture availability that will result in a reduction in deep water, which in turn will reduce shrub density and result in a less conservative nitrogen economy.     

Grazing effects on patchy dryland vegetation in northern Patagonia

In this study the spatial patterns and dynamics of vegetation patches along a grazing gradient in the steppe ofLarrea divaricataandStipaspp. in NE Patagonia (Argentina) are described. A general effect of grazing is the reduction of total plant cover resulting from the decrease in cover of perennial grasses (Stipa speciosa, Poa ligularis, Stipa tenuis) and some tall shrubs (Chuquiraga hystrix, Bougainvillea spinosa, Lycium chilense). Dwarf shrubs (Nassauvia fueguianaandJunellia seriphioides) increase their cover under medium and/or high grazing pressures. Plant species are spatially grouped into patches which alternate with areas of bare soil. Eleven types of vegetation patch differing in the dominant plant functional type or species, floristic richness and size were identified with different relative frequency along the grazing gradient. Based on these results, it is postulated that grazing forces the replacement of large patches dominated by tall shrubs with high species richness, byLarrea divaricatapatches or small dwarf shrub patches with low species richness and the extinction of grass patches. This results from: (1) disruption of local balances of species deletions and additions; (2) fragmentation of large patches; and (3) formation of new vegetation patches. These changes lead to differing plant spatial organization and heterogeneity along the grazing gradient which may be described by characteristic arrays of vegetation patches.     

The influence of elevation, shrub species, and biological soil crust on fertile islands in the Mojave Desert, USA

We quantified soil nutrients and biological crust cover (bryophytes and lichens) under the canopies of three species of Mojave Desert shrubs and in interspaces between shrubs at three elevations to determine the effects of shrub species, soil crust, and elevation on islands of soil fertility. Means of pH, organic matter, total Kjeldahl nitrogen, nitrogen mineralization, and gravimetric soil moisture are significantly greater in soils under Ambrosia dumosa (Gray) Payne, Larrea tridentata Cov., and Coleogyne ramosissima Torr. than soils from adjacent interspace microhabitats. Although soil moisture and soil organic matter increase by a factor of 1.5 from the low elevation to the high elevation site, the ratio of shrub to interspace concentrations, or the difference in mean soil variables between shrubs and interspaces, is effectively constant and independent of elevation. Total bryophyte and lichen cover is relatively low (24.5%), however, there are 11 species of bryophytes and two species of lichens distributed across three elevations with the highest species richness and cover at the low-elevation site. Bryophyte and lichen cover is correlated with silt but is not related, consistently, to soil nutrients. Overall, the balance of processes controlling spatial aggregation of soil nutrients under shrubs is remarkably insensitive to potential differences in organic inputs among elevations, shrub species, and soil crust surfaces.     

Are Fabiana imbricata shrublands advancing over northwestern Patagonian grasslands? A population dynamics study involving fire and precipitation

Shrub encroachment is the advance of woody plants over grasslands. For some shrub species this process is favoured by fire and high precipitation. In northwestern Patagonia spring precipitation is related with the El Niño Southern Oscillation (ENSO). Over the past years, ENSO has shown an increase in both amplitude and frequency. Our study focuses on the population dynamics of the shrub Fabiana imbricata Ruiz & Pav. (Solanaceae) using stochastic models which involve different fire frequencies and variations in spring precipitation. Our results indicate that F. imbricata's population grows at a wide range of fire frequencies, reaching its maximum every 6–10 years. The elasticity analysis show that, as fire frequency diminishes, population growth rate depends almost exclusively on adult survival, recruitment events are rare and the presence of seedlings or juveniles is seldom observed. In this case, the landscape would be formed by mature F. imbricata shrublands, and population persistence would depend on adult survival, seed production and their accumulation in the soil bank. However, in the case of very high fire frequencies, more recurrent recruitment pulses would be observed, with a landscape dominated by young shrublands composed of individuals of different ages and reproductive status.     

Phenology ofMedicago minimaandErodium cicutariumin semi-arid Argentina

The phenology ofMedicago minimavar.minimaandErodium cicutariumwas studied at two different field sites in the Calden District (approx. 10 million ha), a temperate semi-arid phytogeographical region in central Argentina. One site had been protected from wild and domestic herbivores for the 6 previous years. The other adjacent site had been exposed to continuous grazing by cattle for several decades. A phenological key was made for each species in 1988, and this key was improved during the 1989 growing season when phenological observations were made every 1–3 weeks.The growth cycle ofM. minimaandE. cicutariumwas similar at the two studied sites. Initiation of the cycle occurred in autumn and appeared to be associated with soil water availability. Phenological patterns were very dynamic at the reproductive stage in both species. This stage started earlier and was faster inE. cicutariumthan inM. minima. The end of the growing season occurred during late spring, concomitant with high maximum air temperatures. Our results suggest that these species hasten their development when air temperature increases and soil water availability decreases. This could be an important strategy in these species which allows them to persist as seeds, and produce a new generation under favourable environmental conditions.     

The effects of the identity of shrub species on the distribution and diversity of ground arthropods in a sandy desert ecosystem of northwestern China

Shrub is an important factor on structuring ground arthropod communities in desert ecosystems. In this study, in order to determine how shrubs and their species influence ground arthropod distribution patterns in a sandy desert scrubland dominated by two different shrub species, Calligonum mongolicum and Nitraria sphaerocarpa, the ground arthropods were sampled with pitfall traps during spring, summer and autumn. At the community level, total arthropod abundance was shown to be significantly higher under shrubs than in intershrub bare areas in spring; similar patterns occurred in terms of the richness of arthropod groups in the spring and over three seasons, suggesting season-specific shrub presence effects on arthropod activity. In addition, more arthropods were found under N. sphaerocarpa shrubs than under C. rnongolicum shrubs in autumn, suggesting season-specific effects of shrub species of arthropod activity, whereas more arthropods taxa were captured under C. mongoIicum than N. sphaerocarpa. At the trophic group level, the abundances of predator and herbivore arthropods were significantly greater under shrubs than in intershrub bare habitats, whereas herbivore arthropods were more abundant under N. sphaerocarpa than C. rnongolicum, and an opposite rule was detected for predator arthropods At the family level, the mean abundances of Carabidae, Curculionidae, Gnaphosidae and Lycosidae were significantly higher in the shrub microhabitats than in the intershrub bare habitat, there was no significant difference between habitats on the mean abundances of Formicidae and Tenebrionidae. The study results suggested that shrub presence and shrub species variation are important determinants of ground arthropod assemblages in this desert ecosystem, but the responses of ar- thropods differed among trophic and taxonomic groups.     

Influence of shrubs on soil characteristics and their function in Sahelian agro-ecosystems in semi-arid Niger

Soil beneath shrubs form ‘fertile islands’ in fallow sites and millet fields in semi-arid Niger. To gain more information about this phenomena different shrub species in fallow sites following a gradient from 350–650 mm precipitation were examined. For each shrub two different areas were distinguished: an area under the canopy of the shrubs and an area in the nearby open land. Soil samples were taken from a depth of 0–10 cm and analysed for C_org, N_total, P_Bray, pH(H₂O), exchangeable cations, effective cation exchange capacity (ECEC) and soil texture. Significantly higher concentrations between 38–51% for C, N, P and 22% on ECEC for K⁺were found in the soil under the shrubs. The pH showed only slight but significant differences, whereas Al³⁺and H⁺rates on ECEC under the shrubs were increased by 44–55%. For Guiera senegalensis, the most common shrub of the studied area, enrichment ratios of most soil properties increased relatively more with increasing aridity. In general, enrichment ratios decreased with the age of the fallows, whereas concentrations showed no clear evolution. The chemical composition of the shrub litter seems to influence the degree of soil enrichment. The main step of fertile island formation takes place during the cultivation period by trapping wind-blown sediment. This work shows that shrubs are of vital importance for the accumulation of nutrients and maintenance of soil fertility within agro-ecosystems of Niger.     

Climatic response of thick leaf spruce (Picea crassifolia) tree-ring width at different elevations over Qilian Mountains, northwestern China

Tree-ring cores of thick leaf spruce (Picea crassifolia) taken from four sites at different elevations, in the middle of the Qilian Mountains, in the arid and semi-arid region of northwestern China, were used to develop four tree-ring width chronologies using standard dendrochronological methods. Results indicate that with increasing altitude the chronologies’ year-to-year variations decreased. Hence, the sensitivity of the tree-ring chronologies to climate decreases with altitude. Further analysis showed that the significant limiting factor on tree growth is spring precipitation. Measurements of stomata density and leaf dry weight suggest the species’ ecological adaptation strategy changes with elevation. At high elevation the metabolic rate of thick leaf spruce decreases, thus showing the effect of the climate.     

ROOST SITES AND COMMUNAL BEHAVIOR OF ANDEAN CONDORS IN CHILE*

The Andean condor (Vultur gryphus) is an endangered species. Even southern Patagonia, home to the most stable and abundant populations of Andean condors, is witnessing increasing pressure from development and tourism. Taking the case of Torres del Paine National Park, in the Chilean Patagonia, we examine monitoring of condor populations at roosting sites and communal bird behavior in response to humans as an effective tool for bird conservation within protected areas. Based on field data collected throughout 2007, we identify new roosting places, explore activity patterns and population characteristics of free‐ranging and roosting Andean condors, examine bird behavior in response to humans, and analyze the current and likely future ecological impacts of tourism on the condor population and its habitat. Our results reveal that the impact of tourism is still low and that the Andean condors do not seem to be declining in numbers in the park but that the importance of roosts and animal behavior in response to humans must be considered for future monitoring, bird‐conservation planning, and ecotourism management.     

The changing Patagonian landscape: Erosion and westward sediment transfer paths in northern Patagonia during the Middle and Late Pleistocene

Pleistocene glaciations have promoted important landscape transformations as a result of high rates of erosion and rapid sediment evacuation to adjacent marine basins. In the Patagonian Andes the role of the Patagonian Ice Sheet on landscape evolution, in particular the spatial patterns of glacial erosion and its influence on sediment fluxes, is poorly documented. Here, we investigate the Middle and Late Pleistocene sedimentary record of the continental slope from Ocean Drilling Program (ODP) Site 861, offshore Patagonia (46°S), to evaluate the link between glaciations, mountain range erosion and continental margin strata formation. Petrographic analysis of the sand-size fraction (0.063–2 mm) and ɛNd and ⁸⁷Sr/⁸⁶Sr measurements in the silt-size fraction (10–63 µm) indicate that glacial erosion over the last 350,000 years has focused within the Patagonian Batholith, with a minor influence of a proximal source to the drilling site, the Chonos Metamorphic Complex. This shows that erosion has focused in the core of the northern Patagonian Andes, coinciding roughly with the location of the Liquiñe-Ofqui Fault Zone and the zone of concentrated precipitation during glaciations, suggesting a combined climatic and structural control on glacial erosion. Temporal variation in the provenance signal is contemporaneous with a marked change in the stratigraphy of ODP Site 861 that occurred after the glaciation of MIS 8 (~240 kyr ago). Before MIS 8, a restricted provenance signal and coarse lithofacies accumulated on the continental slope indicates spatially restricted erosion and efficient transfer of sediment towards the ocean. In contrast, very high provenance variability and finer continental slope lithofacies accumulation after MIS 8 suggest a disorganized expansion of the areas under erosion and a more distal influence of ice sediment discharge to this site. We argue that this change may have been related to a re-organization of the drainage patterns of the Patagonian Ice Sheet and flow of outlet glaciers to the continental margin during the last two glaciations.     

Structural and nutritional heterogeneity of riparian vegetation in Patagonia (Argentina) in relation to seasonal grazing by sheep

Riparian meadows of southern Patagonia are temporally and spatially heterogeneous habitats. They are thought to play a key role in regulating the dynamics of arid grazed steppes of the region. We conducted a 2-year study with sheep to determine the influence of two grazing conditions (deferment and spring grazing) on structural and nutritional parameters of three vegetation types apparently associated with a soil moisture gradient in a riparian meadow in southern Santa Cruz (Argentina). Spring deferment allowed forage accumulation in very moist and intermediate vegetation types by the beginning of the summer and had no detrimental impact on forage quality. Deferment had no effect on biomass accumulation of the drier vegetation type at our study site. Structural attributes of the vegetation appeared to affect spatial grazing patterns of sheep. Soil-related patchiness, rather than sheep grazing, appeared to control vegetation structure and nutritional value. The proportion of more mesic plant communities in riparian habitats of Patagonia could determine the outcome of plant–animal interactions in these systems. Practical recommendations of grazing capacity in paddocks, or more theoretical considerations of ecosystem dynamics of the Patagonian steppes need to explicitly consider the internal patchiness of riparian habitats.     

Gender-related differences of shrubs in stands of Atriplex canescens with different histories of grazing by cattle

Atriplex canescens is a relatively common dioecious shrub in western North America. It is considered a valuable forage resource for both wild and domestic herbivores. Sex ratios and shrub dimensions were recorded in stands of tetraploid A. canescens that had been either protected from cattle grazing or summer- or winter-grazed by cattle for at least 20 years. Stem diameter and crown shape were used as surrogates for shrub age which could not be estimated by counting growth rings. Shrub sex ratios in exclosures were significantly more female biased than the empirically derived ratio for tetraploid A. canescens (55 female:35 male: 10 monecious). Conversely, shrub sex ratios in grazed pastures were not significantly different from the empirical ratio. Proportion of female shrubs in exclosures was significantly higher than in grazed pastures. Proportion of male shrubs, on the other hand, was similar in exclosures and grazed pastures. Winter-grazed shrub stands were apparently younger than both summer-grazed and protected shrubs. Protected shrubs appeared to be the oldest. Grazed female shrubs were apparently younger than grazed males, however, shrub ages of protected male and female shrubs were apparently not different. Cattle-grazing may have affected female shrubs more negatively at this site, causing gender-based differential mortality, and/or sex-shifting. Such processes could account for the differences in sex ratios, and for the apparent gender-related differences in shrub age that were observed.     

Differences in production and mortality of reproductive structures in two Prosopis L. (Mimosaceae) shrub species from Patagonia, Argentina

Prosopis L. pods have been proposed as a source of gum (galactomannans), with potential applications as a thickening agent in the food industry. The objective of this work was to compare, under field conditions, the production and mortality of reproductive structures in two sympatric shrub species of Prosopis: P. alpataco Phillipi and P. denudans Bentham. Our results indicate that fruit production in Prosopis denudans and P. alpataco is very low and erratic in non-irrigated lands. Although large quantities of flowers (15,000–25,000 flowers*0.40 m⁻²) can be found in one flowering season, 70–80% are shed before they reach anthesis and 20–28% are aborted between anthesis and fruit set. We conclude that the utilization of native populations of Prosopis denudans and P. alpataco for fruit production in Patagonian steppe, could not guarantee a consistent supply for the food industry.     

Stress resistance strategy in an arid land shrub: interactions between developmental instability and fractal dimension

This paper investigates allocation of energy to mechanisms that generate and preserve architectural forms (i.e. developmental stability, complexity of branching patterns) and productivity (growth and reproduction) in response to environmental disturbances (i.e. grazing and resource availability). The statistical error in translational symmetry was used to detect random intra-individual variability during development. This can be thought of as a measure of developmental instability caused by stress. Additionally, we use changes in fractal complexity and shoot distribution of branch structures as an alternate indicator of stress. These methods were applied to Anthyllis cytisoides L., a semi-arid environment shrub, to ascertain the effect of grazing and slope exposure on developmental traits in a 2×2 factorial design. The results show that A. cytisoides maintains developmental stability at the expense of productivity. Anthyllis cytisoides was developmentally more stable when grazed and when on south-facing, as opposed to north-facing slopes. On the contrary, shoot length, leaf area, fractal dimension and reproductive-to-vegetative allocation ratio were larger in north- than in south-facing slopes. As a consequence, under extreme xeric conditions, shrub mortality increased in north-facing slopes, especially when not grazed. The removal of transpiring area and the reduction of plant competition favoured developmental stability and survival in grazed plants. Differences between grazed and ungrazed plants were most evident in more mesic (north-facing) areas.     

Plant water relations of thornscrub shrub species, north-eastern Mexico

As an approach to understand how diurnal and seasonal plant water potentials (Ψ) are related to soil water-content and evaporative demand components, the responses of six thornscrub shrubs species (Havardia pallens, Acacia rigidula, Eysenhardtia texana, Diospyros texana, Randia rhagocarpa, and Bernardia myricaefolia) of the north-eastern region of Mexico to drought stress were investigated during the course of 1 year. All study species showed the typical diurnal pattern of variation in Ψ. That is, Ψ decreased gradually from predawn (Ψ_pd) maximal values to reach minima at midday (Ψ_md) and began to recover in the late afternoon. On a diurnal basis and with adequate soil water-content (>0.20 kg kg⁻¹), diurnal Ψ values differed among shrub species and were negatively and significantly (p<0.001) correlated with air temperature (r=−0.741 to −0.883) and vapor pressure deficit (r=−0.750 to −0.817); in contrast, a positive and significant (p<0.001) relationship was found to exist with relative humidity (r=0.758–0.842). On a seasonal basis, during the wettest period (soil water-content>0.20 kg kg⁻¹), higher Ψ_pd (−0.10 MPa) and Ψ_md (−1.16 MPa) values were observed in R. rhagocarpa, whereas lower figures (−0.26 and −2.73 MPa, respectively) were detected in A. rigidula. On the other hand, during the driest period (soil water-content<0.15 kg kg⁻¹), Ψ_pd and Ψ_md values were below −7.3 MPa; i.e. when shrubs species faced severe water deficit. Soil water-content at different soil layers, monthly mean relative humidity and monthly precipitation were significantly correlated with both Ψ_pd (r=0.538–0.953; p<0.01) and Ψ_md (r=0.431–0.906; p<0.05). Average soil water-content in the 0–50 cm soil depth profile explained between 70% and 87% of the variation in Ψ_pd. Results have shown that when gravimetric soil water-content values were above 0.15 kg kg⁻¹, Ψ_pd values were high and constant; below this threshold value, Ψ declined gradually. Among all shrub species, A. rigidula appeared to be the most drought tolerant of the six species since during dry periods it tends to sustain significantly higher Ψ_pd in relation to B. myricaefolia. The remaining species showed an intermediate pattern. It is concluded that the ability of shrub species to cope with drought stress depends on the pattern of water uptake and the extent to control water loss through the transpirational flux.     

Winter climate change at different temporal scales in Vaccinium myrtillus, an Arctic and alpine dwarf shrub

Snow cover strongly influences plant growth in Arctic and alpine ecosystems. Snow characteristics and snowmelt timing are likely to change in a warmer climate. We studied year rings and shoot growth of the dwarf shrub bilberry ( Vaccinium myrtillus ), and species abundances of the vegetation, in response to early or late snowmelt at a study site in the Central Alps, near Davos, Switzerland. Snowmelt was manipulated on experimental plots for 3 and 30 years. Additional plots were set up along a natural snowmelt gradient, and at high and low elevation. Growth ring data showed an increasing trend in annual growth increment over the last 20 years, especially in the extraordinarily hot summer of 2003. Comparing high and low elevation sites, growth rings were wider at low elevation, but only in cold years. In years with relatively cold summers, however, xylem ring width was greater in plots with late rather than early snowmelt along the natural snowmelt gradient, possibly indicating drought stress in early snowmelt plots. Snow cover had a strong influence on species abundances along the natural snowmelt gradient, and change (not yet significant) was beginning to be seen in plots with 30 years of snow manipulation. Our results indicate that beneficial effects of early snowmelt for shrub growth may be offset in cold summers. Although early snowmelt prolongs the growing season, harsh conditions and frost events early in the growing season may become more likely, and hamper plant growth, and this could affect plant growth in all Arctic and alpine snow-dominated ecosystems.     

Plant Area Index and microclimate underneath shrub species from a Chilean semiarid community

In drylands, environmental conditions under shrub canopy differ from those found in open sites. We should expect that microclimate conditions under shrubs with distinct canopy architecture should also be different. Plant Area Index (PAI) of the three most abundant shrubs species (Porlieria chilensis, Adesmia bedwellii and Proustia cuneifolia) in Bosque Fray Jorge National Park, north-central Chile was measured using a Plant Canopy Analyzer. During two years (2004-2005), we recorded the Relative Humidity and Air Temperature underneath and away from the canopy of the shrubs.The three shrub species showed significant differences in PAI. Microclimate at 30 cm and 2 m above the soil in the open conditions were drier and warmer than underneath shrub canopies. Vegetation patches generate moderate microclimate conditions.Canopy structure can buffer climatic variability, contributing to high herbaceous productivity as well as shrub recruitment. Reflecting shrub architecture and observed PAI values, the lowest microclimate variations were observed under the canopies of P. chilensis, followed by P. cuneifolia and finally A. bedwellii. We bring a novel approach quantifying the Plant Area Index instead of the Plant cover and using a low cost method that integrates the distribution of leaves and may be derived from remote sensing products.     

A Miocene algal assemblage dominated by Pediastrum leonensis n. sp. (Chlorophyceae) from Patagonia, Argentina: paleoenvironmental implications

Taxonomic identification of Pediastrum species in an Early Miocene assemblage from southeast Patagonia, Argentina, is used to assess paleoecological conditions. Pediastrum leonensis n. sp. is described based on unique features of the coenobia morphology. It belongs to the P. kawraiskyi–mustersii–patagonicum complex and has no known living counterpart. This group seems to have a long evolutionary history in the region. P. kawraiskyi and P. mustersii are recorded for the first time in pre-Quaternary deposits and their presence suggests a depositional environment represented by a cold stenothermal lake conditions, indicating a temperate to a cold-temperate climate.     

Annual growth zones in stems of Hypericum irazuense (Guttiferae) in the Costa Rican páramos

The high peaks of the Cordillera de Talamanca in Costa Rica support shrub- and grass-dominated páramo ecosystems that experience stand-replacing wildfires. The dry season that facilitates these fires results in dormancy in plant growth and provides an opportunity to use dendrochronological analyses to determine ages of plants in burn sites to support studies of fire history and postfire vegetation recovery. This study investigates the formation of annual growth zones in stems of the common shrub Hypericum irazuense. Unlike other páramo shrubs, H. irazuense rarely resprouts following fire, instead recovering through seedling recruitment following seed dispersal from the unburned periphery. Laboratory analysis of 19 prepared cross sections from 15 stems shows that samples of H. irazuense from burned areas can provide an estimate of the minimum number of years since the previous fire, supporting earlier work based on field examination of stems. Including a time lag for seedling recruitment or resprouting refines that estimate. Counts of growth zones in most sections coincided with dates of the last known fires. The presence of annual growth zones in H. irazuense places the species within a relatively small group of woody Neotropical species for which annual rings or growth zones have been demonstrated.