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Main traits related to fire adaptation in Pinus canariensis . a) Thick bark, insulating the cambium from high temperatures in a 30-year-old individual; bar = 10 cm. b) and c) Epicormic resprouting after crown scorching by fire from preformed, preventitious buds; bar in b) = 5 cm. d) Closed, xeriscent cones in a 45 year-old pine. Only serotinous trees show this profuse cone-bearing, otherwise scarce in this species.
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A wide set of phenotypic characteristics related to life history were studied in mature stands of Pinus canariensis throughout its natural range of distribution in the Canary archipelago. Natural forests ranging from those located in xeric
areas through to the sub-tropical cloud forests and high mountain stands were classified into eight ecological...
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Citations
... Los incendios forestales son considerados el principal agente perturbador de este hábitat a largo plazo (Climent et al., 2004). Estos han sido recurrentes en el pinar canario desde la llegada del ser humano al archipiélago (Arévalo et al., 2001;Gil y González Doncel, 2013) y, en la actualidad, su frecuencia e intensidad se han visto favorecidas por acción del cambio climático . ...
... The Canary pine forest was particularly affected due to the location of the new volcano along the slope of Cumbre Vieja. Pinus canariensis has been influenced by volcanic eruptions for millions of years (Anderson et al., 2009) and exhibits many traits consistent with adaptations to fire (Climent et al., 2004) and volcanic eruptions (López de Heredia et al., 2014). It can withstand heavy metal pollution associated with volcanic eruptions (Rodríguez Martín et al., 2013), resprouts from all Overview of the 100 sample locations on the island of La Palma (black crosses). ...
Volcanic eruptions have a strong environmental impact on surrounding forests. Trees are affected by mechanical damage, tephra deposition and volcanic gases. Oceanic islands are shaped by relatively frequent volcanic eruptions and thus offer the opportunity to study the effect of volcanic activity on biodiversity. We investigate the impact of volcanic gas emissions and tephra deposition during the 2021 Tajogaite eruption on the Canary Pine forests of the island of La Palma, Spain, characterized by monospecific stands of the endemic pine species Pinus canariensis C. Sm. ex D.C. Large quantities of volcanic sulphur dioxide caused chlorotic damage up to approximately 7 km around the crater, followed by widespread resprouting of P. canariensis.
To detect the spatial pattern of impacts, we sampled P. canariensis needles from all over the island of La Palma and analyzed their sulphur (S), nitrogen (N) and carbon (C) content. We found a strong increase of S needle content close to the crater, while C decreased significantly. S levels were strongly related to distance to the crater, C and N were mostly influenced by S content. Trees affected by volcanic gases allocate resources to resprouting, leading to lower levels of C due to translocation of C as a building block. Surprisingly, we found higher N levels in needles with high levels of S and a less clear pattern compared to C, likely due to a multitude of environmental factors influencing N needle levels.
We investigated how canopy damage patterns detected in Sentinel-2 remote sensing imagery after the eruption correlated to the in-situ needle contents. However, we did not find a clear correlation between in-situ needle values and spectral responses in remote sensing. While satellite images were well suited to analyse large scale patterns of canopy damage following the eruption, needle levels varied strongly on a local, tree-based level, which is not reflected in remote sensing imagery.
... intrusion of lowland species adapted to warmer climates into cooler communities; Duque, Stevenson & Feeley, 2015;Fadrique et al., 2018) of montane forests in the Caribbean region (Tanner et al., 2022). Increased fire incidence linked to dryer atmospheric conditions could facilitate the expansion of fire-resistant species (Grau & Veblen, 2000), for instance, serotinous pine trees (Rodríguez-Trejo & Fulé, 2003;Climent et al., 2004). In some regions (e.g. ...
In spite of their small global area and restricted distributions, tropical montane forests (TMFs) are biodiversity hotspots and important ecosystem services providers, but are also highly vulnerable to climate change. To protect and preserve these ecosystems better, it is crucial to inform the design and implementation of conservation policies with the best available scientific evidence, and to identify knowledge gaps and future research needs. We conducted a systematic review and an appraisal of evidence quality to assess the impacts of climate change on TMFs. We identified several skews and shortcomings. Experimental study designs with controls and long-term (≥10 years) data sets provide the most reliable evidence, but were rare and gave an incomplete understanding of climate change impacts on TMFs. Most studies were based on predictive modelling approaches, short-term (<10 years) and cross-sectional study designs. Although these methods provide moderate to circumstantial evidence, they can advance our understanding on climate change effects. Current evidence suggests that increasing temperatures and rising cloud levels have caused distributional shifts (mainly upslope) of montane biota, leading to alterations in biodiversity and ecological functions. Neotropical TMFs were the best studied, thus the knowledge derived there can serve as a proxy for climate change responses in under-studied regions elsewhere. Most studies focused on vascular plants, birds, amphibians and insects, with other taxonomic groups poorly represented. Most ecological studies were conducted at species or community levels, with a marked paucity of genetic studies, limiting understanding of the adaptive capacity of TMF biota. We thus highlight the long-term need to widen the methodological, thematic and geographical scope of studies on TMFs under climate change to address these uncertainties. In the short term, however, in-depth research in well-studied regions and advances in computer modelling approaches offer the most reliable sources of information for expeditious conservation action for these threatened forests.
... One of the few conifers which is considered as a good resprouter in both juvenile and adult stages is the Canary Island pine (Pinus canariensis C.Sm. ex DC; Pausas and Keely 2017). In this species, the resprouting capability has been proposed to be linked to its evolutionary history in a volcanic environment under selective pressures exerted by perturbation regimes and driving the dynamics of its populations (Climent et al. 2004). Moreover, small injuries promote the cessation of cambial activity in the wounded area (Chano et al. 2015), including the repression of genes involved in radial growth (Chano et al. 2017a, b). ...
Resprouting is crucial in population dynamics in response to wildfires or herbivory. This trait, common in angiosperms, is considered rare in conifers, being Pinus canariensis one of the few species able to resprout. We analyzed gene expression during wound-induced resprouting in 5 years-old Canarian pines. RNA was extracted at three different stages, including control samples from remote branches, representing immediate response to wounding (R0), resprouting initiation (R1), and elongation (R2), and then hybridized to a microarray designed with 15,000 cDNAs from Pinus canariensis transcriptome of meristematic activity. We found 1164 Differentially Expressed Genes (DEGs), of which 903 were significant in R0, 278 in R1, and 261 in R2. Genes related to defense- and stress-response were mainly found to be overexpressed at R0, while DEGs coding for transcription factors such as GROWTH-REGULATING FACTOR (GRF), FLOWERING-PROMOTING FACTOR (FPF), and the HOMEODOMAIN LEUCINE ZIPPER Class IV (HD-ZIP IV), mainly related to outgrowth processes and lateral organogenesis in plants, showed overexpression at R1 and R2, when new shoots were emerging. This data was compared to differential gene expression during apical growth in P. canariensis , suggesting similarities between vegetative apical growth and regulation of lateral shoot development in response to mechanical wounding, but also yielding 32 DEGs exclusively overexpressed at some point during resprouting stages (R1 and/or R2). The implication of such transcription factor families, with important roles in leaf and inflorescence development also in flowering plants, suggests underlying homologies between common lateral organogenesis processes in angiosperms and resprouting in gymnosperms.
... In the highlands of Gran Canaria there are currently patches of "pristine" stands of pine forests (Pinus canariensis) (Pérez de Paz et al., 1994). The Canarian pine forest is considered to be adapted to high-intensity fires with traits such as thick bark insulating cambium, epicormic resprouting, and serotinous cones, suggesting a certain degree of resilience to recurrent disturbance by natural fires (Climent et al., 2004;Keeley and Zedler, 1998). Ravazzi et al. (2021) found that before the island was settled, the pine forest dominating the summit was already subjected to recurrent fires. ...
Islands contribute enormously to global biodiversity, but are threatened by human activities. The emergence of island societies has transformed island ecosystems since prehistory. Focusing on drivers of change related to human activities, such as land(sea)-use change, resource extraction, pollution, invasive and alien species, and climate change, we revise the history of human impacts on islands with special focus on species extirpation ofs and ecosystem changes.
... The Canary Island pine was thus notably affected by tephra fall, sulphuric acid aerosol 12 , and short episodes of acid rain. However, this conifer shows high resistance to temperature, confirming its great adaptation to volcanic events 16 , which is probably also one of the keys to its resistance to the more frequent present-day wildfires 17 . This pine species has evolved among volcanoes for the last 13 My 16 and has adapted successfully to high temperatures. ...
Volcanic activity provides a unique opportunity to study the ecological responses of organisms to catastrophic environmental destruction as an essential driver of biodiversity change on islands. However, despite this great scientific interest, no study of the biodiversity at an erupting volcano has yet been undertaken. On La Palma (Canary archipelago), we quantified the main species affected and their fate during the 85-day eruption (September–December 2021). Our main objective consisted of monitoring the biodiversity subjected to critical stress during this volcanic eruption. We found that all biodiversity within a 2.5 km radius was severely affected after the first two weeks. It is challenging to assess whether volcanism can drive evolutionary traits of insular organisms. Examples are the adaptation of an endemic conifer to high temperatures, selection of functional plant types—secondary woodiness—, effects of the disappearance of invertebrates and their influence in trophic nets and vertebrate trophic plasticity. However, our data suggest that such previous evolutionary changes might continue to favour their resilience during this eruption. Lastly, it is a very good opportunity to assess the extent to which these periodic volcanic catastrophes may constitute temporary windows of repeated opportunities for the evolution and speciation of oceanic island biota.
... Species can have other fire adaptation than bark thickness, like sprouting and serotiny. Q. suber has a corky bark as well as its ability to sprout, while P. canariensis has serotiny in addition to thick corky bark (Climent et al., 2004). Because plant employ several adaptational fire resistance means, the relationship between bark thickness and fire resistance may not always strong. ...
Pines (Pinus, with 120 species) and oaks (Quercus, with at least 400 species), the two principal forest forming genera of the Northern Hemisphere occur in fire affected landscapes over a large geographical area from 120°W to 150°E long, and from 10° to 60°N lat. We compiled studies of 116 such sites of which 75 sites had data of bark thickness. Taking a meta-analysis approach, here we have compared the absolute and relative bark thickness (bark thickness per unit stem diameter) of oak species sharing mixed stands to generalize adaptational significance of bark in relation to fire. For the meta-analysis we considered only natural forests with adequate evidences of fire history. We gave importance to relative bark thickness, which is less sensitive to tree size than absolute thickness while comparing species. At 100 cm girth the average absolute bark thickness was significantly higher for Diploxylon pines (3.65 cm) than for oaks (2.2 cm) and Haploxylon pines (1.65 cm). On an average, the relative bark thickness was higher for the fire adapted Diploxylon pines (6.3%) than for Quercus species, (4.4%). The resource allocation efficiency was higher in pine species than for oak species, emphasizing the importance of bark characters in their persistence and range expansion. The difference between species of two principal forest genera in relative bark thickness and hence success in fire affected landscapes across a wide geographical range strongly testifies its adaptational significance to fire.
... In Pinus, development of juvenile-like shoots after juvenile-adult phase transition is infrequent, but it can take place. For example, occurrence of epicormic juvenile-like shoots has been reported in P. canariensis and P. rigida after fire damage (Climent et al., 2004;Gucker, 2007), after trimming in young trees of P. echinata (Bormann, 1955), and in mature trees of P. maximartinezii (Moreno-Letelier and Vázquez-Lobo, personal observation). Therefore, the incidence of juvenilelike shoots in the adult vegetative phase of conifers indicates that this developmental pattern in some cases could be activated under environmental stress. ...
In land plants, heteroblasty broadly refers to a drastic change in morphology during growth through ontogeny. Juniperus flaccida and Pinus cembroides are conifers of independent lineages known to exhibit leaf heteroblasty between the juvenile and adult life stage of development. Juvenile leaves of P. cembroides develop spirally on the main stem and appear decurrent, flattened, and needle‐like; whereas adult photosynthetic leaves are triangular or semi‐circular needle‐like, and grow in whorls on secondary or tertiary compact dwarf shoots. By comparison, J. flaccida juvenile leaves are decurrent and needle‐like, and adult leaves are compact, short, and scale‐like. Comparative analyses were performed to evaluate differences in anatomy and gene expression patterns between developmental phases in both species. RNA from 12 samples was sequenced and analyzed with available software. They were assembled de novo from the RNA‐Seq reads. Following assembly, 63,741 high‐quality transcripts were functionally annotated in P. cembroides and 69,448 in J. flaccida. Evaluation of the orthologous groups yielded 4140 shared gene families among the four references (adult and juvenile from each species). Activities related to cell division and development were more abundant in juveniles than adults in P. cembroides, and more abundant in adults than juveniles in J. flaccida. Overall, there were 509 up‐regulated and 81 down‐regulated genes in the juvenile condition of P. cembroides and 14 up‐regulated and 22 down‐regulated genes in J. flaccida. Gene interaction network analysis showed evidence of co‐expression and co‐localization of up‐regulated genes involved in cell wall and cuticle formation, development, and phenylpropanoid pathway, in juvenile P. cembroides leaves. Whereas in J. flaccida, differential expression and gene interaction patterns were detected in genes involved in photosynthesis and chloroplast biogenesis. Although J. flaccida and P. cembroides both exhibit leaf heteroblastic development, little overlap was detected, and unique genes and pathways were highlighted in this study. Juniperus flaccida and Pinus cembroides exhibit leaf heteroblasty between the juvenile and adult life stage of development. Comparative analyses were performed to evaluate differences in anatomy and gene expression patterns between developmental phases in both species.
... Within a few years, the previous canopy cover can be reached again [49]. The species additionally has very thick bark and a high concentration of carbohydrates in the sapwood parenchyma cells [47,50]. The extremely long needles of this pine aid substantially in freshwater provision [51], by contributing to condensation during the frequent fog (clouds touching the ground). ...
In autumn 2021, the largest volcanic eruption on the island of La Palma in historic records took place. The Canary Islands are of volcanic origin and eruptions have always constituted part of their natural disturbance regime. Until recently, their impacts could not be directly observed and studied. Influence of the emission of phytotoxic gases on biodiversity and ecosystem dynamics was hitherto unknown. The recent eruption is still being intensely monitored. We used Sentinel-2 remote sensing data to analyze the spatial extent and intensity of the impact related to sulfuric emissions, aiming to understand the damage patterns in Canary pine forest. The emissions damaged 10% of that forest and affected 5.3% of the Natura 2000 protected areas. We concluded that this is largely due to the toxic effects of the enormous emissions of SO2. We found a clear correlation between the change in the normalized difference vegetation index (NDVI) and distance from the eruption. This pattern was weakly anisotropic, with stronger damage in southern directions. Counteracting effects, such as ash deposition, were largely excluded by combining NDVI change detection with tree cover density. We expect that vegetation damage will be transient. P. canariensis can resprout after forest fires, where most leaves are lost. Consequently, our assessment can serve as a reference for future ecosystem regeneration.
... Today, anthropogenic causes of wildfires, whether deliberate or accidental, add to the natural drivers; only a small proportion of the fires on the Canary Islands is now attributed to ignition by natural causes [19,22]. Fire frequency may have been even higher in the last few centuries, when agriculture was the main economic activity. ...
... The Canary pine forest ecosystem differs from many other fire-prone forest types. P. canariensis is well adapted to frequent and intense wildfires [22]. It has thick, protective bark, serotinous cones and readily re-sprouts from all above-ground organs and roots after fire. ...
... P. canariensis forests play a key role in maintaining the water balance of several of the Canary Islands. In the canopies of these pine trees, the moisture from clouds condenses on their long pine needles, significantly increasing the total precipitation inside the forest on the windward eastern slopes [22]. Due to the constant trade winds being blocked by the island's topography, this cloud contact is very frequent. ...
Forest fires are drivers of spatial patterns and temporal dynamics of vegetation and biodiversity. On the Canary Islands, large areas of pine forest exist, dominated by the endemic Canary Island pine, Pinus canariensis C. Sm. These mostly natural forests experience wildfires frequently. P. canariensis is well-adapted to such impacts and has the ability to re-sprout from both stems and branches. In recent decades, however, anthropogenically caused fires have increased, and climate change further enhances the likelihood of large forest fires. Through its dense, long needles, P. canariensis promotes cloud precipitation, which is an important ecosystem service for the freshwater supply of islands such as La Palma. Thus, it is important to understand the regeneration and vegetation dynamics of these ecosystems after fire. Here, we investigated species diversity patterns in the understory vegetation of P. canariensis forests after the large 2016 fire on the southern slopes of La Palma. We analyzed the effect of fire intensity, derived from Sentinel-2 NDVI differences, and of environmental variables, on species richness (alpha diversity) and compositional dissimilarity (beta diversity). We used redundancy analysis (dbRDA), Bray–Curtis dissimilarity, and variance partitioning for this analysis. Fire intensity accounted for a relatively small proportion of variation in alpha and beta diversity, while elevation was the most important predictor. Our results also reveal the important role of the endemic Lotus campylocladus ssp. hillebrandii (Christ) Sandral & D.D.Sokoloff for understory diversity after fire. Its dominance likely reduces the ability of other species to establish by taking up nutrients and water and by shading the ground. The mid- to long-term effects are unclear since Lotus is an important nitrogen fixer in P. canariensis forests and can reduce post-fire soil erosion on steep slopes.
