The Scientific World Journal

The Scientific World Journal / 2014 / Article

Research Article | Open Access

Volume 2014 |Article ID 215418 | 16 pages |

Prescribed Burning and Clear-Cutting Effects on Understory Vegetation in a Pinus canariensis Stand (Gran Canaria)

Academic Editor: Regis Cereghino
Received13 Feb 2014
Revised23 Jun 2014
Accepted07 Jul 2014
Published24 Jul 2014


Prescribed fires are a powerful tool for reducing fire hazards by decreasing amounts of fuel. The main objective is to analyze the effects of prescribed burning on the understory vegetation composition as well as on the soil characteristics of a reforested stand of Pinus canariensis. The study attempts to identify the effects of the preburning treatment of cutting understory vegetation on the floristic parameters of the vegetation community. This study was carried out for two years following a prescribed fire in a Canarian pine stand. Cutting and burning treatment affected species composition and increased diversity. Burnt and cut plots were characterized by a diverse array of herbaceous species and by a lower abundance of Teline microphylla (endemic legume), although burning apparently induced its germination. Cut treatment was more consistently differentiated from the control plots than burnt treatment. Soil K decreased after both treatments, pH slightly decreased after cutting, while P and Ca increased after fire. From an ecological point of view, prescribed burning is a better management practice than cutting the woody species of the understory. However, long-term studies would be necessary to evaluate the effects of fire intensity, season and frequency in which the prescribed burning is applied.

1. Introduction

Historically, fire has played a dominant role in shaping many forest plant communities [1]. Mediterranean-type vegetation is one of the world’s major fire-prone biomes [2], with conifer forests among the most flammable ecosystems in the Mediterranean region [3]. In these ecosystems, fire is a crucial process controlling vegetation dynamics and structure [4]. Species have developed different strategies to survive after fire, whether at the individual level, with a thick insulating bark or the ability to resprout from underground parts, or at the population level, with serotinous cones or seeds which are resistant to or stimulated by high temperatures.

Fire regimes have changed as a consequence of human activities and large wildland fires are now more likely to occur. Changing socioenvironmental conditions, such as abandonment of traditional agricultural crops, abandonment of pastoralism, or decreasing exploitation of timber resources, are leading to higher fuel loads and consequently to the increase in the frequency and severity of wildfires [5, 6]. As a result, wildfire prevention measures are necessary and prescribed fires are a powerful management tool towards this goal [1]. In particular, low-intensity prescribed fires are considered useful in some ecosystems to reduce fuel loads by decreasing the amount of low vegetation and small decayed wood [7, 8] without important effects on soil properties [9].

The most recent studies on the effects of prescribed fire in Europe are related to shrublands [4, 10] or grasslands [9], but few deal with pine forests [11]. Studies on vegetation dynamics after wildfires may be used to assess the possible responses of vegetation to prescribed fire under similar conditions. Most studies indicate that although herb and some shrub cover are strongly affected [7], species abundance [1215] and diversity [16, 17] increase following fire.

Regarding soil properties, prescribed burning has been shown to increase pH and nutrient availability immediately following fire and through the first year [9, 18, 19]. However, these changes usually revert to preburn values after one year. In several studies carried out on the Canarian pine forest, this effect on nutrient content seems to be more persistent [2025], being possible to detect from 1 up to even 17 years after fire for some nitrogen and phosphorous parameters [2224]. In contrast, in case of some parameters, such as pH or exchangeable cations, differences might disappear between several months and two years after fire [25, 26].

Canarian forest stands have been subjected to long-term degradation, especially since the European colonization of the islands [27]. Despite being the largest forest community of the islands, only 54% of the Canarian pine forest (60,678 ha) persists nowadays in its natural extension [28]. Restoration programs of the Canarian pine forest have been very common since 1930, existing 15,103 ha of Canarian pine plantation, which require management practices to reach a more natural stage [29]. In this framework, prescribed burning can operate not only as a management tool for fire prevention, but also as a tool for achieving restoration objectives [30].

Little is known about historic fire regimes in the Canary Islands, which makes it difficult to determine which conditions and frequency of burning would be the most appropriate to restore vegetation as well as prevent large wildfires. In the past, wildfire were infrequent but large in extension [20], but after human settlement the frequency of fires increased [33]. At present, natural fire regime has been clearly exceeded, especially in recent decades; thus, most of the Canarian pine forests have been burned in the last 25–30 years, and several stands in that time frame have done so two or three times [34].

If fire management is based on a misunderstanding of plant life history or incorrect historical perceptions, burning could have potentially large effects on forest community diversity [35]. In particular, Pinus canariensis is specifically adapted to fire [36], displaying both resistant and resilient strategies, which evidently makes the role of fire during the evolution of this ecosystem important. Moreover, the Canarian pine forest is an ecosystem of particular interest since it has a high percentage of endemic flora and is distinct from other Mediterranean ones. Thus, it is important to study the effect of fire management on this particular ecosystem.

Management of pine forest in Mediterranean countries usually involves cutting the shrub understory to break fuel continuity within the canopy [14]. This practice is also used in the Canary Islands and thus we differentiated between two treatments, understory cutting and subsequent prescribed burning (hereafter “burnt”) and only preburning management (hereafter “cut”). Burning without cutting was not possible due to the danger of canopy torching under the present fuel model. Cutting reduces understory cover and the abundance of vertical fuel ladders. However, due to debris in the field after the cutting treatment, fuel loads remain high and light availability to the soil surface is reduced, which can also affect species composition.

The objective of this study was to analyze the effects of clearing and prescribed fire treatments on the understory community and soil parameters by testing the following main hypotheses: (1) cutting and burning will have different effects on species composition, and different species will be identified as descriptors of each treatment; (2) burning will modify soil parameters more intensively than just cutting, but some of these parameters such as pH or exchangeable cations will recover to pretreatment values in less than two years.

2. Methods

2.1. Study Site

The study was conducted at the pine forest of Artenara, in Gran Canaria, Canary Islands, Spain (UTM-X 436449, UTM-Y 3099622), which is part of the Protected Landscape of Las Cumbres, under environmental protection by the Canarian Network of Natural Protected Areas [37]. This is a young forest, about 60 years old, which has hardly been managed and never burned since it was planted during restoration management. Pine density is approximately 600 trees per ha with high fuel accumulation. The study site is located between 1400 and 1600 m a.s.l. and faces the prevailing north easterly winds. Mean annual temperature, humidity, and rainfall (for 2006–2008) are 17.7°C, 52.2%, and 500 mm, respectively. The dominant tree species is Pinus canariensis, although the area includes other planted exotic pine species such as P. halepensis and P. radiata. The understory vegetation is dominated by a variety of shrub and herb species, where the most representative species are Chamaecytisus proliferus, Teline microphylla, and Micromeria benthami. A deep litter layer, with an average thickness of 5.7 cm, typically covers the entire site.

2.2. Design of the Experiment

The study site was divided into six plots of between 1.5 and 2 ha each, depending on topographical complexity, where experimental treatments (3 cut and 3 burnt) were applied, and three control plots of approximately 0.5–1 ha each. Control plots were smaller than treated plots to meet management objectives in the largest possible area. From several weeks to a month before burning, the woody understory vegetation of all plots and the lowest branches of the pines were cleared, with the help of chainsaws, except in the controls, as in usual practice to break the vertical continuous fuel ladder before burning. Dead fuel was kept on the ground. This practice allows a homogeneous burning and maintains flame heights of less than 1.5 m due to the spreading of the dead fuel along the surface covering fuel gaps [38]. Three of these six cleared plots were chosen at random and burnt in June 2006 (burnt treatment) while in the other tree plots they were keep as cut treatment. Burning was carried out under specific temperature (18–26°C), humidity (30–70%), and wind (<15 km/h) conditions. Strip fires, a burn method that consists in setting successive parallel strips of fire, and back fires, in which the line of fire is set on the upslope side of the fuel and the fire moves slowly against the wind and slope, were used to achieve ignition. Based on all these parameters and that the canopy only was reached by the flames occasionally, the burning was considered of medium intensity.

Three 100 m2 square subplots per treatment plot and one per control plot were located at random. Environmental variables such as rock and litter cover percentage were visually estimated per subplot. Altitude, aspect, and slope were also measured per subplot. All the species in the subplots were listed and their percentage cover was visually estimated and noted on a scale of 1 to 9 (1: trace; 2: <1%; 3: 1-2%; 4: 2–5%; 5: 5–10%; 6: 10–25%; 7: 25–50%; 8: 50–75%; 9: >75%) [31]. Vegetation was sampled six, twelve, eighteen, and twenty-four months after burning.

Four samples of the top 10 cm soil (only organic horizon) were taken per subplot and pooled to obtain a single composite sample before analysis. Soil samples were analyzed following the common standard methods for organic carbon (Walkey-Black method), available phosphorus (Olsen method), potassium, magnesium, calcium, and sodium (Bower method) [39]. Soil pH was analyzed following extraction with deionised water and measured with a pH-meter.

Vegetation and soil sampling were carried out before treatment (spring 2005) and six (winter 2006), twelve (spring 2007; only vegetation), eighteen (winter 2007), and twenty-four (spring 2008) months after burning.

2.3. Statistical Analysis

To ensure that there were no significant differences in understory species composition, richness, and diversity before burning and cutting treatment, a distance based permutational MANOVA [40] was fitted, with treatment area (control, burnt, and cut) as fixed factor. The analyses were based on Bray-Curtis distances of the cover data of vascular species and on Euclidian distances of the number of species and diversity prior to treatment (spring 2005). A maximum of 9999 permutations were used to obtain the values (). Shannon diversity index: , where is the proportion of species relative to the total number of species, was used to characterize species diversity of the pine stand.

In order to test our hypothesis of species composition changes after management practices, distance based permutational-repeated measures MANOVA was fitted, with treatment (control, burnt, and cut) and period (four repeated measures: six, twelve, eighteen, and twenty-four months after treatment) used as fixed factors and the plots as a paired factor. The analyses were based on Bray-Curtis distances of the cover data of understory vascular species. Same procedure was applied to species richness and diversity data but based on Euclidian distances. Significant terms were investigated using a posteriori pairwise comparisons with the PERMANOVA t-statistic. A maximum of 9999 permutations were used to obtain the values () and the Monte Carlo correction was applied where necessary.

We performed a similarity percentage (SIMPER) routine [41], using the Bray-Curtis coefficient, to identify both, descriptor species associated with each treatment and those taxa that primarily contribute to average dissimilarity between treatments [42]. This was accompanied by a principal components analysis (PCA) to graphically represent variation in species composition, as it helps to reveal environmental variables not included in the sampling method [43]. We introduced understory species cover data as variables and one datum per treatment per sampling period (the information of the nine subplots that composed the treatment which were pooled into only one, assuming the average of the abundance for the species, based on understory cover data) as samples to reinforce general patterns of species composition depending on the treatment and for a better understanding of the graph. No covariable was introduced for the analysis as none of the environmental variables measured was significantly different between treatments. Primer 6 and PERMANOVA + (PRIMER-E Ltd., Plymouth, UK) were used to perform all statistical procedures, with the exception of the PCA, which was performed using CANOCO for Windows [44].

Finally, to test our second hypothesis we performed one-way distance based permutational ANOVA [40], comparing soil parameters between treatments (fixed factor) for each time period. Analyses were based on Euclidian distances. Significant terms were investigated using a posteriori pairwise comparisons with the PERMANOVA t-statistic. A maximum of 9999 permutations were used to obtain the values () in each data set and the Monte Carlo correction was applied where necessary.

3. Results

3.1. Descriptors and Species Composition

The absence of significant differences in species composition (Pseudo-, ), richness (Pseudo-, ), and diversity (Pseudo-, ) between the treatment areas before being cut and burnt provides assurance that the differences detected between them in the following sampling period are result of management treatments.

We found a total of 107 species belonging to 39 families (12 species were only identified at genus level) and 44% of the species were common to the different treatment areas (Table 4). Cutting and burning treatment affected species composition and diversity ( SD;  SD;  SD) but not species richness (Table 1), which was on average  SD. Absence of interaction between treatment and sampling period indicates that changes are maintained over time, as can be appreciated in the PCA graph (Figure 1), where the samples are clearly separated along axis I, with control samples located on the left side of the graph, and treated samples on the right. This separation is mostly due to the abundance of subshrubs and woody species (e.g., Teline microphylla, Sonchus acaulis, Erysimum bicolor, Artemisia thuscula, and Argyranthemum adauctum) being more abundant in the control plots, together with some herbaceous species (e.g., Ranunculus cortusifolius, Sonchus oleraceus, Todaroa montana, and Galium aparine). These species play consistent roles (mean dissimilarity to SD ratio higher than 1) in determining the dissimilarity between treatments, especially between control and cut plots more separated in the bidimensional space of the PCA. Micromeria benthamii was more abundant in cut than in control plots, while Hirschfeldia incana and Lathyrus annuus were more abundant in burnt than in control plots (SIMPER; Table 5). Cut and burnt plots were separated along axis II, being characterized mainly by a diverse array of herbaceous species (Figure 1). Although, according to the SIMPER procedure (Table 5) only a few species consistently discriminated between the two treatments. Based on both SIMPER (average dissimilarity between treatments: control versus cut = 72.35, control versus burnt = 61.52, and cut versus burnt = 72.06) and PCA results the cut treatment was more consistently differentiated from the control plots than burnt treatment (Figure 1; Table 5).

Species compositionSpecies richnessSpecies diversity

PERMANOVAPseudo- (perm)Pseudo- (perm)Pseudo- (perm)

Sampling period1.590.021.380.240.340.91
Treatment * sampling period0.680.850.270.950.420.88

Pairwise -test (perm) (perm) (perm)


Regarding descriptive species responsible for the similarity within treatments, presented using the results from the SIMPER procedure (Table 2), only few species play more or less consistent roles in determining the similarity within treatments. T. microphylla was the dominant species in all treatments, having the largest abundance in control plots. As shown above E. bicolor and S. acaulis were also abundant and consistently present within control plots (Table 2). These three species account for more than 45% of the similarity within control plots. This pattern changed after the treatments, with different results in cut and burnt plots. Cut plots were very variable, without any descriptive species especially abundant and/or consistent within the treated plots. In contrast, burnt plots were more homogeneous, including T. microphylla (especially abundant as seedlings) and the herb Silene vulgaris as descriptive species (Table 2).

SpeciesAv. Abund.Av. Sim.Sim./SDContrib. %Cum. %

Control (average similarity 54.59)
Teline microphylla (DC.) P. E. Gibbs & Dingwall6.2514.252.5526.1126.11
Erysimum bicolor (Hornem.) DC.2.676.523.3311.9438.05
Sonchus acaulis Dum. Cours.2.754.741.368.6946.74
Silene vulgaris (Moench) Garcke2.084.552.098.3355.07
Sonchus oleraceus L.1.834.534.678.3163.38
Ranunculus cortusifolius Willd.2.424.301.367.8971.26
Todaroa montana Webb ex Christ2.082.700.824.9476.20
Galium aparine L.1.332.460.974.5080.70
Artemisia thuscula Cav.1.251.380.642.5383.23
Hirschfeldia incana (L.) Lagr.-Foss.
Ferula linkii Webb1.171.230.532.2687.82
Argyranthemum adauctum (Link) Humphries1.001.160.532.1389.94
Salvia canariensis L.1.080.960.311.7591.70

Cut (average similarity 26.30)
Teline microphylla (DC.) P. E. Gibbs & Dingwall1.645.120.7419.4819.48
Silene vulgaris (Moench) Garcke1.534.770.7618.1337.61
Micromeria benthamii Webb & Berthel.1.423.950.6815.0152.63
Vicia disperma DC.0.611.840.217.0059.63
Erysimum bicolor (Hornem.) DC.
Sonchus acaulis Dum. Cours.0.890.980.413.7268.03
Galium aparine L.0.830.920.443.4971.52
Todaroa montana Webb ex Christ0.920.810.423.0874.60
Lathyrus annuus L.0.690.750.382.8677.46
Ranunculus cortusifolius Willd.1.000.660.392.5279.98
Sonchus oleraceus L.0.690.620.342.3582.33
Hirschfeldia incana (L.) Lagr.-Foss.0.670.520.331.9884.31
Adenocarpus foliolosus (Aiton) DC.0.690.440.161.6685.97
Stellaria media (L.) Vill.0.640.420.301.5987.56
Argyranthemum adauctum (Link) Humphries0.750.400.301.5489.09
Echium onosmifolium Webb0.360.360.171.3790.46

Burnt (average similarity 37.33)
Teline microphylla (DC.) P. E. Gibbs & Dingwall2.117.202.3919.2719.27
Silene vulgaris (Moench) Garcke2.006.161.6116.4935.76
Sonchus acaulis Dum. Cours.1.562.840.717.6143.37
Erysimum bicolor (Hornem.) DC.1.582.790.707.4850.85
Sonchus oleraceus L.1.252.650.987.0957.94
Hirschfeldia incana (L.) Lagr.-Foss.1.392.330.826.2364.17
Ranunculus cortusifolius Willd.1.422.100.675.6369.81
Lathyrus annuus L.0.921.300.543.4973.30
Galium aparine L.0.780.830.442.2475.53
Chamaecytisus proliferus (L. f.) Link0.860.800.382.1577.69
Todaroa montana Webb ex Christ0.940.780.302.1079.79
Centranthus calcitrapae (L.) Dufr.0.690.650.371.7581.53
Andryala pinnatifida Aiton0.780.650.411.7583.28
Argyranthemum adauctum (Link) Humphries0.750.600.391.6284.90
Echium onosmifolium Webb1.390.570.281.5486.43
Papaver rhoeas L.0.640.490.341.3087.73
Galium parisiense L.0.670.480.331.2989.02
Stellaria media (L.) Vill.0.560.430.341.1690.18

3.2. Soil Parameters

The absence of significant differences for any of the soil variables prior to treatment (Table 3) provides assurance that the differences detected between them in the following sampling periods are result of management treatments.

Prefire6 months18 months24 months

pH6.27 (0.15)6.29 (0.20)6.46 (0.18)6.5 (0.15)a6.18 (0.24)b6.74 (0.23)a6.2 (0.75)6.28 (0.37)6.52 (0.32)6.47 (0.50)6.44 (0.24)6.51 (0.31)
% OM3.47 (0.25)4.04 (0.67)3.95 (0.72)5.03 (2.03)5.28 (1.42)4.98 (1.89)3.87 (1.06)4.26 (0.82)4.03 (0.74)4.27 (0.96)4.5 (0.60)4.08 (1.01)
ppm P12.67 (4.62)11.56 (5.73)12.75 (5.75)10.67 (3.06)a8.89 (7.75)a21.11 (8.25)b14 (5.29)ab10 (5.39)a19.78 (7.97)b6.67 (4.16)7.56 (7.73)12.67 (7.42)
Ca9.33 (1.70)8.58 (1.50)9.75 (1.48)9.53 (2.37)ab8.82 (1.87)a11.82 (1.94)b11.13 (3.79)9.31 (2.08)12.02 (3.70)11 (3.89)9.38 (1.78)10.76 (3.24)
Mg5.53 (1.70)4.47 (1.50)4.83 (1.48)4.47 (2.37)4.09 (1.87)4.18 (1.94)5.53 (3.79)4.47 (2.08)5 (3.70)6.47 (3.89)4.82 (1.78)4.84 (3.24)
K1.37 (1.70)1.29 (1.50)1.3 (1.48)2.3 (2.37)a1.64 (1.87)b1.7 (1.94)b1.9 (3.79)1.5 (2.08)1.56 (3.70)2.00 (3.89)1.47 (1.78)1.43 (3.24)
Na0.87 (0.15)0.91 (0.20)0.93 (0.18)1.37 (0.15)1.21 (0.24)1.32 (0.23)2.5 (0.75)2.18 (0.37)2.12 (0.32)1.83 (0.50)1.51 (0.24)1.4 (0.31)

C: control; CT: cut; B: burnt; different letters indicate significant differences (permutational ANOVA, 9999 permutations, ; pairwise -test, ).


AlliaceaeAllium sp.All_sp.xx
ApiaceaeFerula linkii WebbFer_linESxxx
ApiaceaeTodaroa montana Webb ex ChristTod_monEGxxx
Apiaceae Torilis nodosa (L.) Gaertn. Tor_nodxx
AsteraceaeAndryala pinnatifida AitonAnd_pinESxxx
AsteraceaeArgyranthemum adauctum (Link) HumphriesArg_adaESxxx
AsteraceaeArtemisia thuscula Cav.Art_thuESxxx
AsteraceaeCalendula arvensis L.Cal_arvMNxx
AsteraceaeCarduus pycnocephalus L.Car_pycMNxxx
AsteraceaeCarduus tenuiflorus CurtisCar_tenMNx
AsteraceaeCentaurea aspera L.Cen_aspMNxxx
AsteraceaeConyza sp.Con_sp.SIx
AsteraceaeGalactites tomentosus MoenchGal_tomMNxxx
AsteraceaeHedypnois rhagadioloides (L.) F. W. SchmidtHed_rhaMNx
AsteraceaeHypochaeris glabra L. Hyp_glaMNxxx
AsteraceaeLactuca serriola L. Lac_serMNxx
AsteraceaeLeontodon taraxacoides (Vill.) Mérat Leo_tarPIx
AsteraceaePericallis webbii Sch. Bip. & BollePer_webESxxx
AsteraceaeReichardia tingitana (L.) Roth Rei_tinPNxx
AsteraceaeSenecio teneriffae Sch. Bip.Sen_tenSNx
AsteraceaeSonchus acaulis Dum. Cours.Son_acaESxxx
AsteraceaeSonchus asper (L.) A. W. Hill Son_aspMNxxx
AsteraceaeSonchus oleraceus L.Son_oleMNxxx
AsteraceaeTolpis barbata (L.) Gaertn.Tol_barMNxx
AsteraceaeTragopogon porrifolius L.Tra_porMNx
AsteraceaeUrospermum picroides (L.) Scop. ex F. W. SchmidtUro_picPNx
BoraginaceaeEchium onosmifolium WebbEch_onoESxxx
Boraginaceae Myosotis
BrassicaceaeErysimum bicolor (Hornem.) DC.Ery_bicSNxxx
BrassicaceaeHirschfeldia incana (L.) Lagr.-Foss.Hir_incMNxxx
BrassicaceaeRaphanus raphanistrum L.Rap_rapSNxx
BrassicaceaeSisymbrium officinale (L.) Scop.Sis_offMNx
CampanulaceaeLegousia hybrida (L.) Delarbre Leg_hybMNxxx
CampanulaceaeWahlenbergia lobelioides (L. f.) Link Wah_lobSNx
CaryophyllaceaePinus radiata D. Don Pin_radSIxx
CaryophyllaceaeSilene gallica L.Sil_galMNxx
CaryophyllaceaeSilene vulgaris (Moench) GarckeSil_vulMNxxx
CaryophyllaceaeStellaria media (L.) Vill.Ste_medPIxxx
Caryophyllaceae Petrorhagia nanteuilii (Burnat) P. W. Ball & Heywood Pet_nanx
CaryophyllaceaeCerastium glomeratum Thuill. Cer_gloMNxxx
ChenopodiaceaeChenopodium sp.Che_sp.x
CrassulaceaeAeonium percarneum (R. P. Murray) Pit.Aeo_perESxxx
CrassulaceaeAeonium simsii (Sweet) StearnAeo_simESxx
CrassulaceaeGreenovia aurea (C. Sm. ex Hornem.) Webb & Berthel.Gre_aurEGxx
CrassulaceaeUmbilicus gaditanus Boiss.Umb_gadPNxxx
DipsacaceaePterocephalus dumetorus (Brouss. ex Willd.) Coult.Pte_dumESxx
EricaceaeErica arborea L.Eri_arbSNx
FabaceaeAdenocarpus foliolosus (Aiton) DC.Ade_folESx
FabaceaeChamaecytisus proliferus (L. f.) LinkCha_proESxxx
FabaceaeLathyrus annuus L.Lat_annPIxxx
FabaceaeTeline microphylla (DC.) P. E. Gibbs & DingwallTel_micESxxx
Fabaceae Trifolium arvense L. Tri_arvPNxx
FabaceaeTrifolium campestre Schreb. in SturmTri_camPNxxx
FabaceaeTrifolium scabrum L.Tri_scaPNxxx
Fabaceae Trifolium
FabaceaeVicia disperma DC.Vic_disPNxxx
FabaceaeVicia lutea L.Vic_lutMNxxx
Fabaceae Vicia sativa L. Vic_satMNxx
Fabaceae Vicia
Fumariaceae Fumaria
GeraniaceaeErodium cicutarium (L.) L`Hér. in AitonEro_cicMNxx
GeraniaceaeGeranium dissectum L. Ger_disMNx
GeraniaceaeGeranium molle L.Ger_molMNxx
GeraniaceaeGeranium purpureum Vill.Ger_purMNxx
GeraniaceaeGeranium rotundifolium L. Ger_rotMNxx
HypericaceaeHypericum reflexum L. f.Hyp_refESx
Hypolepidaceae Pteridium aquilinum (L.) Kuhn in Kerst. Pte_aquPNx
IridaceaeRomulea columnae Sebast. & MauriRom_colSNxx
LamiaceaeLamium amplexicaule L. Lam_ampSIxx
LamiaceaeMicromeria benthamii Webb & Berthel.Mic_benESxxx
LamiaceaeSalvia canariensis L.Sal_canESxxx
LamiaceaeStachys arvensis (L.) L. Sta_arvMNxx
OrchidaceaeNeotinea maculata (Desf.) StearnNeo_macPNxxx
Oxalidaceae Oxalis pes-caprae L. Oxa_pesIIx
PapaveraceaePapaver rhoeas L.Pap_rhoIIxxx
PinaceaePinus canariensis C. Sm. ex DC. in BuchPin_canESxxx
Pinaceae Pinus halepensis Mill. Pin_halSIx
Pinaceae Polycarpon tetraphyllum (L.) L.Pol_tetMNx
PoaceaeAira caryophyllea L. Air_carMNxx
PoaceaeAnisantha madritensis (L.) NevskiAni_madMNx
PoaceaeAnisantha rigida (Roth) Hyl.Ani_rigMNx
PoaceaeAvena barbata Pott ex LinkAve_barMNxx
PoaceaeAvena fatua L.Ave_fatSNx
PoaceaeBriza maxima L.Bri_maxMNxx
PoaceaeCynosurus echinatus L.Cyn_echMNx
PoaceaeLamarckia aurea (L.) MoenchLam_aurPNx
PoaceaeVulpia geniculata (L.) LinkVul_genMNxx
PolygonaceaeRumex bucephalophorus L.Rum_bucSNxxx
PolygonaceaeRumex lunaria L.Rum_lunESx
PrimulaceaeAsterolinon linum-stellatum (L.) DubyAst_linPNxxx
RanunculaceaeRanunculus cortusifolius Willd.Ran_corSNxxx
ResedaceaeReseda luteola L. Res_lutPNx
RubiaceaeGalium aparine L.Gal_apaMNxxx
RubiaceaeGalium parisiense L.Gal_parPNxxx
RubiaceaeSherardia arvensis L.She_arvMNxx
Rubiaceae Galium murale (L.) All. Gal_murPNx
Rubiaceae Galium sp.Gal_sp.x
Rubiaceae Galium verrucosum Huds. Gal_verMNx
ScrophulariaceaeMisopates orontium (L.) Raf. Mis_oroPNx
ScrophulariaceaeVeronica hederifolia L.Ver_hedMNxx
SolanaceaeSolanum nigrum L. Sol_nigMNxx
UrticaceaeForsskaolea angustifolia Retz.For_angESx
UrticaceaeUrtica sp.Urt_sp.x
ValerianaceaeCentranthus calcitrapae (L.) Dufr.Cen_calMNxxx

SpeciesControlCutAv. Diss.Diss./SDContrib. %Cum. %
Av. Abund.Av. Abund.

Teline microphylla (DC.) P. E. Gibbs & Dingwall6.251.648.951.6112.3712.37
Sonchus acaulis Dum. Cours.2.750.894.001.345.5317.90
Erysimum bicolor (Hornem.) DC.2.671.223.761.205.2023.10
Ranunculus cortusifolius Willd.2.421.003.721.355.1428.24
Todaroa montana Webb ex Christ2.080.923.301.104.5632.80
Sonchus oleraceus L.1.830.692.551.373.5236.33
Salvia canariensis L.
Chamaecytisus proliferus (L. f.) Link0.670.722.430.563.3643.07
Micromeria benthamii Webb & Berthel.0.171.422.381.143.2846.35
Galium aparine L.1.330.832.101.022.9149.26
Artemisia thuscula Cav.
Ferula linkii Webb1.
Silene vulgaris (Moench) Garcke2.081.531.940.872.6957.69
Argyranthemum adauctum (Link) Humphries1.000.751.921.012.6660.35
Hirschfeldia incana (L.) Lagr.-Foss.1.000.671.741.132.4062.76
Stellaria media (L.) Vill.0.830.641.620.942.2565.00
Echium onosmifolium Webb0.750.361.560.752.1567.15
Andryala pinnatifida Aiton0.750.561.480.912.0569.20
Vicia disperma DC.0.330.611.450.712.0171.21
Lathyrus annuus L.0.330.691.220.961.6972.90
Pericallis webbii Sch. Bip. & Bolle0.670.111.180.771.6374.53
Umbilicus gaditanus Boiss.0.670.191.170.731.6176.14
Adenocarpus foliolosus (Aiton) DC.0.000.691.140.451.5877.73
Asterolinon linum-stellatum (L.) Duby0.500.311.090.761.5179.24
Papaver rhoeas L.0.580.141.040.811.4480.68
Centranthus calcitrapae (L.) Dufr.0.330.361.010.571.4082.08
Rumex bucephalophorus L.0.250.560.940.631.3083.38
Hypochaeris glabra L. 0.250.390.890.601.2384.60
Carduus pycnocephalus L.0.080.470.700.590.9785.57
Aeonium percarneum (R. P. Murray) Pit.0.330.190.690.600.9686.53
Trifolium scabrum L.
Aeonium simsii (Sweet) Stearn0.420.000.660.530.9188.37
Galium parisiense L.0.330.080.640.470.8989.26
Neotinea maculata (Desf.) Stearn0.170.190.630.440.8790.13

 SpeciesControlBurntAv. Diss.Diss./SDContrib. %Cum. %
Av. Abund.Av. Abund.

Teline microphylla (DC.) P. E. Gibbs & Dingwall6.252.116.751.7510.9810.98
Todaroa montana Webb ex Christ2.080.943.081.175.0115.99
Sonchus acaulis Dum. Cours.2.751.563.011.264.9020.89
Ranunculus cortusifolius Willd.2.421.422.731.274.4425.33
Erysimum bicolor (Hornem.) DC.2.671.582.431.123.9529.28
Echium onosmifolium Webb0.751.392.180.883.5532.83
Chamaecytisus proliferus (L. f.) Link0.670.862.090.713.4036.23
Salvia canariensis L.1.080.332.090.713.4039.62
Ferula linkii Webb1.170.691.990.953.2342.85
Artemisia thuscula Cav.1.250.501.831.062.9745.83
Galium aparine L.1.330.781.771.112.8748.70
Hirschfeldia incana (L.) Lagr.-Foss.1.001.391.741.162.8351.53
Argyranthemum adauctum (Link) Humphries1.000.751.560.952.5454.07
Andryala pinnatifida Aiton0.750.781.410.982.2956.36
Stellaria media (L.) Vill.0.830.561.390.982.2558.62
Lathyrus annuus L.0.330.921.371.072.2360.85
Sonchus oleraceus L.1.831.251.310.932.1262.97
Centranthus calcitrapae (L.) Dufr.0.330.691.220.781.9864.95
Papaver rhoeas L.0.580.641.220.991.9866.93
Silene vulgaris (Moench) Garcke2.
Umbilicus gaditanus Boiss.0.670.441.180.791.9270.76
Galium parisiense L.0.330.671.070.761.7572.51
Pericallis webbii Sch. Bip. & Bolle0.670.031.000.761.6374.14
Carduus pycnocephalus L.0.080.720.950.711.5575.69
Vicia disperma DC.0.330.440.950.641.5577.24
Micromeria benthamii Webb & Berthel.0.170.440.840.551.3678.60
Asterolinon linum-stellatum (L.) Duby0.500.060.830.671.3679.96
Stachys arvensis (L.) L. 0.000.530.810.591.3181.27
Trifolium scabrum L.0.250.420.730.661.1882.45
Aeonium simsii (Sweet) Stearn0.420.220.720.631.1783.62
Hypochaeris glabra L.
Vicia lutea L.0.170.330.580.510.9485.58
Rumex bucephalophorus L.
Aeonium percarneum (R. P. Murray) Pit.0.330.110.550.570.8987.37
Trifolium campestre Schreb. in Sturm0.080.360.540.550.8888.25
Geranium rotundifolium L. 0.000.390.480.560.7789.02
Erica arborea L.0.000.310.470.330.7789.79
Lactuca serriola L. 0.000.390.470.500.7690.55

SpeciesCutBurntAv. Diss.Diss./SDContrib. %Cum. %
Av. Abund.Av. Abund.

Erysimum bicolor (Hornem.) DC.1.221.583.410.974.734.73
Sonchus acaulis Dum. Cours.0.891.563.180.874.419.14
Ranunculus cortusifolius Willd.1.001.422.961.004.1013.25
Micromeria benthamii Webb & Berthel.1.420.442.880.914.0017.25
Todaroa montana Webb ex Christ0.920.942.700.763.7521.00
Hirschfeldia incana (L.) Lagr.-Foss.0.671.392.701.003.7524.75
Chamaecytisus proliferus (L. f.) Link0.720.862.550.763.5428.28
Echium onosmifolium Webb0.361.392.490.743.4531.73
Sonchus oleraceus L.0.691.252.471.033.4335.16
Silene vulgaris (Moench) Garcke1.532.002.470.763.4338.59
Teline microphylla (DC.) P. E. Gibbs & Dingwall1.642.112.360.773.2741.86
Lathyrus annuus L.0.690.922.130.902.9544.82
Vicia disperma DC.0.610.441.970.642.7447.55
Galium aparine L.0.830.781.950.892.7050.26
Argyranthemum adauctum (Link) Humphries0.750.751.730.882.4052.66
Andryala pinnatifida Aiton0.560.781.670.872.3254.98
Centranthus calcitrapae (L.) Dufr.0.360.691.550.752.1657.14
Stellaria media (L.) Vill.0.640.561.550.812.1559.28
Carduus pycnocephalus L.0.470.721.500.742.0961.37
Adenocarpus foliolosus (Aiton) DC.0.690.001.400.421.9463.31
Ferula linkii Webb0.060.691.370.461.9065.21
Stachys arvensis (L.) L. 0.220.531.370.581.8967.11
Papaver rhoeas L.0.140.641.200.661.6768.78
Galium parisiense L.0.080.671.190.631.6570.43
Geranium rotundifolium L. 0.190.391.040.541.4471.87
Trifolium scabrum L.0.280.420.980.561.3673.23
Hypochaeris glabra L. 0.390.190.970.561.3574.58
Rumex bucephalophorus L.0.560.220.920.591.2875.86
Umbilicus gaditanus Boiss.0.190.440.900.561.2577.11
Salvia canariensis L.0.220.330.890.461.2378.34
Artemisia thuscula Cav.0.060.500.840.601.1679.50
Lactuca serriola L. 0.110.390.780.531.0880.57
Trifolium campestre Schreb. in Sturm0.080.360.760.471.0681.63
Erodium cicutarium (L.) L’Hér. in Aiton0.360.220.680.550.9482.57
Silene gallica L.0.250.360.670.570.9383.50
Erica arborea L.0.000.310.670.320.9284.42
Vicia lutea L.0.060.330.630.410.8785.29
Hypericum reflexum L. f.
Asterolinon linum-stellatum (L.) Duby0.310.060.530.440.7486.79
Pterocephalus dumetorus (Brouss. ex Willd.) Coult.
Cerastium glomeratum Thuill.
Neotinea maculata (Desf.) Stearn0.190.000.480.320.6688.81
Lamium amplexicaule L.
Oxalis pes-caprae L.
Solanum nigrum L.

We only detected significant influence of the treatment on some soil parameters six and eighteen months after treatment. Six months after treatment, mean pH values were significantly different between treatments (Pseudo-, ), being lower in cleared plots than in control and burnt ones (Table 3). However, pH recovered to original values 18 months after treatments. P concentrations were significantly different between treatments six (Pseudo-, ) and eighteen months after fire (Pseudo-, ), when P concentrations were higher in burnt plots. Although eighteen months after fire these differences only persist between burnt and cut plots (Table 3). Ca content was significantly lower in cut plots than in burnt ones six months after fire (Pseudo-, ). Finally, K was significantly lower in treated than in control plots six months after management treatment (Pseudo-, ).

4. Discussion

Prescribed fire is a useful management tool for preventing large wildfires [1] but can have significant effects on vegetation community diversity [35] and species composition [45], depending on how and where it is applied. Most sources suggest an overall trend of increasing species richness with prescribed fire [13, 15, 16, 45] and a rapid recovery after a low-to-moderate-intensity fire [13, 26]. In most cases, these increases in richness take place during the first to second year after fire and are related to herbaceous pioneer species [15, 45, 46], being linked to canopy opening and higher light availability at soil surface after fire [47]. But studied treatments, understory clearing (cut) and low-intensity prescribed fire (cut), do not imply canopy opening, explaining the absence of significant differences in species richness between treatments (Table 1).

Contrary to species richness, the abundance of understory plants is more sensitive to management [48], leading to the detected changes in species composition (Table 1; Figure 1). The main species leading to differences between treatments was T. microphylla, being more abundant in control plots and explaining more than 10% of the dissimilarities found with management treatments. Studies on the soil seed bank suggest that T. microphylla germination is stimulated by fire [49], which agree with the results of this research, since despite its lower abundance T. microphylla was identified as a descriptor species in burnt plots, where it was especially abundant as seedling but not in cut plots.

Legume shrubs typical of the understory of the pine forest, such as Adenocarpus foliolosus and Chamaecytisus proliferus, can be classified as pyrophitic species, showing a fast recovery or even increasing their cover after a wildfire [21, 26, 50, 51]. In this study C. proliferus was the most abundant in burnt plots, coinciding with recent studies in natural pine forest of the island [26]. Although this species did not consistently discriminate between treatments, it was among the ten species that contributed most to explain the dissimilarities between control and burnt plots (Table 5). In addition, it has been proven that herbaceous legumes play an important role during the early years after fire, showing a typical pioneer strategy of fast growth and propagule production [14, 26, 52]. Consistent with this, L. annuus was more abundant in burnt than in control plots, as were Vicia spp. and Trifolium spp. (Table 5).

Species composition has changed due to management, and two years after the treatment differences remain (Table 1, Figure 1). These differences focus mainly on the higher abundance of some shrubs and perennial herbs in control plots. However attending to the cover classes, with exception of T. microphylla, these differences are of small magnitude, remaining in the range of 1-2% of cover. Consequently, we are assuming that only a few more years will be necessary for the shrubs and herbaceous species to increase their abundance in the treated plots, as has been found in other Canarian pine forest stands where only a few years are necessary to recover the prefire species composition [26].

As established in our first hypothesis, both treatments revealed different effects on species composition. Some authors found that, in high-density pine stands, minimal disturbances, such as surface fires, have a low impact on understory composition and production [50, 53]. Fire is an intrinsic element of the Canarian pine forest [54], which can favour regeneration of several understory species [55], which could explain that in the studied pine stand based on SIMPER and PCA results the cut treatment was more consistently differentiated from the control plots than burnt treatment. From this point of view, fire appears as a disturbance for which adaptation is complete in the plant community and offers a natural heterogenic landscape that favours diversity as has been found in other studies, while cutting leads to higher differences with the original forest stand and to lower species diversity (Table 1) [50].

The impact of prescribed fire on soil nutrients is known to be more evident during the first year after fire, increasing the levels of the most important elements (P, N, K, and Ca), while in general after one year, soil nutrient content is more similar between burned and nonburned plots [9, 18, 19]. However, in the Canarian pine forest, fire effects on soil properties seem to be more persistent [2024]. According to the literature, wildfire effects on nitrogen and phosphorous might be evident 17 years after fire [2224], changes in pH can be maximum 2-3 years after fire [20], and organic matter might need more than four years to recover prefire values [26]. Nonetheless, long lasting effects of fire on soil nutrients are not a rule in Canarian pine forest, since some studies also reveal that pH and exchangeable cations recover prefire values in less than two years [25, 26], and these short-term effects of fire in soil nutrient have also been found in this study.

As hypothesized, fire modified soil parameters more intensively than just cutting, and treatment effects on soil variables lasted less than two years. Burning effects on soil nutrient can differ in direction and intensity from those caused by clearing treatments [55]. Forest management through clear-cutting can result in a pH decrease [56], as has been found in this study (Table 3), probably as a result of the acidification effects of the pine needles that remained as debris in the field after the cutting treatment. On the other hand, fire impact and ash deposition can increase soil pH [20, 55, 57], which usually returns to prefire values within a short period [19, 20]. However, significant increases might occur only at high temperatures [57], which might have not been generated by the low-intensity prescribed fire experienced in this study, explaining the unexpected absence of fire effects on this variable.

Increases in P and exchangeable cations in the surface soil after a low-intensity prescribed fire have been found as common [9, 19, 5860], due to the combustion of organic matter and the heating conversion of nutrients from nonavailable to available forms [57, 61, 62]. Although the amount of P was doubled as a result of prescribed fire, this positive effect was not long lasting, as found in other studies [57]. Despite different effects of burning and clearing on soil nutrients [55, 56], detected difference in Ca content between these two treatments might have arisen due to a tendency to lower content of this element in cut plots, already present in initial soil conditions (Table 1). For soil K, a slight decrease, compared to control plots, was found six months after both treatments, probably because this mobile cation might have been lost through leaching [55]. Absence of fire impact on some of the analyzed soil parameters or discrepancies regarding intensity, direction, and durability of these effects with previous studies [2226] could be the result of differences in fire severity, in burned understory species, and on initial soil nutrient content [57, 6062].

4.1. Management Implications

Based on the obtained results, from an ecological point of view, prescribed fire appears to be a better management practice than simply cutting the woody understory. On the one hand, although none of the management practices resulted in pernicious effects on soil nutrient content, prescribed burning seems to favour a short-term pulse in soil nutrients. On the other hand, two years after treatments fuel reduction aims are accomplished, since dominant shrub species (T. microphylla) abundance remains in the range of 1-2% of cover in treated plots, while in control plots average abundance range is between 10 and 25% (Table 2). Nevertheless, after two years, higher species composition similarity between control and burnt plots indicates that, in comparison to clearing, prescribed burning accelerates autosuccession back to prefire vegetation conditions.

It should be noted that the stimulation of T. microphylla in burnt plots may be counterproductive if it exceeds pretreatment biomass; nevertheless, longer-term studies are necessary to see how this species develops and how long it takes to attain prefire conditions and exceed them, if this is the case, as well as the evaluation of fire intensity, the season, and the frequency in which the prescribed burning is applied.

Perhaps being an efficient and cheap method of forest fuel removal and fire hazard reduction [63, 64], as well as a tool for increasing recreational values and restoring fire prone ecosystems [32, 65], active management through prescribed burning is very limited and rather unpopular in the Canary Islands and in other regions of Spain [64]. However we believe that, in order to avoid catastrophic fires that endanger property and even human lives (e.g., the recent fires on Gran Canaria in 2013 and La Gomera and Tenerife in 2012), a change in fire management should be implemented, encouraging an active management through prescribed burning, since from an ecological point Canarian pine forest has the ability to recover from fire effects in a short term [26].

Conflict of Interests

The authors declare that there is no conflict of interests regarding the publication of this paper.


The authors thank Consejería de Medio Ambiente y Emergencias (Gran Canaria Council), especially UOFF and PRESA, for granting permission to work in Pinus canariensis forest, carrying out the prescribed fire, and partially supporting this study (CI02380503). They also thank the Ministerio de Educación y Ciencia (MEC) of the Spanish Government, which provided financial support by granting C. García a FPU (Formación de Profesorado Universitario, AP2005-4736) predoctoral fellowship. Many thanks are to the nice staff of Gran Canaria Council and to Lea de Nascimento, Tamar de la Concepción, Oscar Socas, and students of the Ecology Department (University of La Laguna) for their help in the fieldwork. The authors would especially like to thank Marcos Salas for his help in plant identification.


  1. R. J. Whelan, The Ecology of Fire, Cambridge University Press, Cambridge, UK, 1995.
  2. Z. Naveh, “The evolutionary significance of fire in the Mediterranean region,” Vegetatio, vol. 29, pp. 199–208, 1975. View at: Google Scholar
  3. H. R. Chandle, P. Cheney, P. Thomas, L. Trabaud, and D. Williams, Fire in Forestry Volume I: Forest Fire Behaviour and Effects, Wiley, New York, NY, USA, 1983.
  4. M. J. Baeza, A. Valdecantos, J. A. Alloza, and V. R. Vallejo, “Human disturbance and environmental factors as drivers of long-term post-fire regeneration patterns in Mediterranean forests,” Journal of Vegetation Science, vol. 18, no. 2, pp. 243–252, 2007. View at: Publisher Site | Google Scholar
  5. D. M. Molina-Terrén, F. Grillo, and D. García-Marco, “Uso del fuego prescrito para la creación de rodales cortafuegos: estudio del caso “Las mesas de Ana López”, Vega de San Mateo, Gran Canaria, España,” Investigación Agraria: Sistemas y Recursos Forestales, vol. 15, pp. 271–276, 2006. View at: Google Scholar
  6. F. Moreira, O. Viedma, M. Arianoutsou et al., “Landscape—wildfire interactions in southern Europe: implications for landscape management,” Journal of Environmental Management, vol. 92, no. 10, pp. 2389–2402, 2011. View at: Publisher Site | Google Scholar
  7. R. J. Phillips and T. A. Waldrop, “Changes in vegetation structure and composition in response to fuel reduction treatments in the South Carolina Piedmont,” Forest Ecology and Management, vol. 255, no. 8-9, pp. 3107–3116, 2008. View at: Publisher Site | Google Scholar
  8. A. Youngblood, C. S. Wright, R. D. Ottmar, and J. D. McIver, “Changes in fuelbed characteristics and resulting fire potentials after fuel reduction treatments in dry forests of the Blue Mountains, Northeastern Oregon,” Forest Ecology and Management, vol. 255, no. 8-9, pp. 3151–3169, 2008. View at: Publisher Site | Google Scholar
  9. X. Úbeda, M. Lorca, L. R. Outeiro, S. Bernia, and M. Castellnou, “Effects of prescribed fire on soil quality in Mediterranean grassland (Prades Mountains, north-east Spain),” International Journal of Wildland Fire, vol. 14, no. 4, pp. 379–384, 2005. View at: Publisher Site | Google Scholar
  10. M. de Luis, J. Raventós, and J. C. González-Hidalgo, “Post-fire vegetation succession in Mediterranean gorse shrublands,” Acta Oecologica, vol. 30, no. 1, pp. 54–61, 2006. View at: Publisher Site | Google Scholar
  11. F. Moreira, A. Delgado, S. Ferreira et al., “Effects of prescribed fire on vegetation structure and breeding birds in young Pinus pinaster stands of northern Portugal,” Forest Ecology and Management, vol. 184, no. 1–3, pp. 225–237, 2003. View at: Publisher Site | Google Scholar
  12. L. Calvo, R. Tárrega, and E. Luis, “Regeneration in Quercus pyrenaica ecosystems after surface fires,” International Journal of Wildland Fire, vol. 1, pp. 205–210, 1991. View at: Publisher Site | Google Scholar
  13. E. E. Knapp, D. W. Schwilk, J. M. Kane, and J. E. Keeley, “Role of burning season on initial understory vegetation response to prescribed fire in a mixed conifer forest,” Canadian Journal of Forest Research, vol. 37, no. 1, pp. 11–22, 2007. View at: Publisher Site | Google Scholar
  14. B. Pérez and J. M. Moreno, “Fire-type and forestry management effects on the early postfire vegetation dynamics of a Pinus pinaster woodland,” Plant Ecology, vol. 134, no. 1, pp. 27–41, 1998. View at: Publisher Site | Google Scholar
  15. F. C. Rego, S. C. Bunting, and J. M. da Silva, “Changes in understory vegetation following prescribed fire in maritime pine forests,” Forest Ecology and Management, vol. 41, no. 1-2, pp. 21–31, 1991. View at: Publisher Site | Google Scholar
  16. L. Trabaud and J. Lepart, “Diversity and stability in garrigue ecosystems after fire,” Vegetatio, vol. 43, no. 1-2, pp. 49–57, 1980. View at: Publisher Site | Google Scholar
  17. L. Calvo, S. Santalla, E. Marcos, L. Valbuena, R. Tárrega, and E. Luis, “Regeneration after wildfire in communities dominated by Pinus pinaster, an obligate seeder, and in others dominated by Quercus pyrenaica, a typical resprouter,” Forest Ecology and Management, vol. 184, no. 1–3, pp. 209–223, 2003. View at: Publisher Site | Google Scholar
  18. P. Kutiel and Z. Naveh, “The effect of fire on nutrients in a pine forest soil,” Plant and Soil, vol. 104, no. 2, pp. 269–274, 1987. View at: Publisher Site | Google Scholar
  19. B. M. Rau, J. C. Chambers, R. R. Blank, and D. W. Johnson, “Prescribed fire, soil, and plants: burn effects and interactions in the central Great Basin,” Rangeland Ecology & Management, vol. 61, no. 2, pp. 169–181, 2008. View at: Publisher Site | Google Scholar
  20. P. Höllermann, “The impact of fire in Canarian ecosystems 1983–1998,” Erdkunde, vol. 54, no. 1, pp. 70–75, 2000. View at: Publisher Site | Google Scholar
  21. J. S. Notario, D. M. Afonso, and R. Díaz, “Effect of a wildfire on volcanic soils under pine forest and broom scrub in Tenerife (Canary Islands),” in Proceedings of the 13th International Soil Conservation Organization Conference, Brisbane, Australia, paper 739, 2004. View at: Google Scholar
  22. J. Durán, A. Rodríguez, J. M. Fernández-Palacios, and A. Gallardo, “Changes in soil N and P availability in a Pinus canariensis fire chronosequence,” Forest Ecology and Management, vol. 256, pp. 384–387, 2009. View at: Google Scholar
  23. J. Durán, A. Rodríguez, J. M. Fernández-Palacios, and A. Gallardo, “Long-term decrease of organic and inorganic nitrogen concentrations due to pine forest wildfire,” Annals of Forest Science, vol. 67, no. 2, p. 207, 2010. View at: Publisher Site | Google Scholar
  24. A. Rodríguez, J. Durán, J. M. Fernández-Palacios, and A. Gallardo, “Wildfire changes the spatial pattern of soil nutrient availability in Pinus canariensis forests,” Annals of Forest Science, vol. 66, no. 2, pp. 210p1–210p7, 2009. View at: Publisher Site | Google Scholar
  25. A. Hernández, N. Rodríguez, J. Notario, C. D. Arbelo, and A. Rodríguez-Rodríguez, “Evaluation of changes in soil in the short and medium term due to a forest fire in the pine forest of Tenerife (Canary Islands, Spain),” FLAMMA, vol. 4, no. 3, pp. 166–170, 2013. View at: Google Scholar
  26. J. A. Arévalo, S. Fernández-Lugo, A. Naranjo-Cigala et al., “Post-fire recovery of an endemic Canarian pine forest,” International Journal of Wildland Fire, vol. 23, no. 3, pp. 403–409, 2014. View at: Publisher Site | Google Scholar
  27. J. J. Parsons, “Human influences on the pine and Laurel Forests of the Canary Islands,” Geographical Review, vol. 71, pp. 253–271, 1981. View at: Google Scholar
  28. M. del Arco, R. González-González, V. Garzón-Machado, and B. Pizarro-Hernández, “Actual and potential natural vegetation on the Canary Islands and its conservation status,” Biodiversity and Conservation, vol. 19, pp. 3089–3140, 2010. View at: Google Scholar
  29. J. R. Arévalo and J. M. Fernández-Palacios, “From pine plantations to natural stands. Ecological restoration of a Pinus canariensis sweet, ex spreng forest,” Plant Ecology, vol. 181, no. 2, pp. 217–226, 2005. View at: Publisher Site | Google Scholar
  30. C. Messier, K. J. Puettmann, and K. D. Coates, Managing Forests as Complex Adaptive Systems: Building Resilience to the Challenge of Global Change, Earthscan from Routledge, 2013.
  31. E. van der Maabel, “Transformation of cover-abundance values in phytosociology and its effects on community similarity,” Vegetatio, vol. 39, no. 2, pp. 97–114, 1979. View at: Publisher Site | Google Scholar
  32. M. Arechavaleta, S. Rodríguez, N. Zurita, and A. García, Lista de especies silvestres de Canarias. Hongos plantas y animales terrestres 2009, Gobierno de Canarias, Islas Canarias, Spain, 2010.
  33. S. Nogué, L. de Nascimento, J. M. Fernández-Palacios, R. J. Whittaker, and K. J. Willis, “The ancient forests of La Gomera, Canary Islands, and their sensitivity to environmental change,” Journal of Ecology, vol. 101, no. 2, pp. 368–377, 2013. View at: Publisher Site | Google Scholar
  34. J. R. Arévalo and J. M. Fernández-Palacios, “9550 Pinares endémicos canaries,” in VV.AA. Bases Ecológicas Preliminares Para la Conservación de los Tipos de Hábitat de Interés Comunitario en España, Dirección General de Medio Natural y Política Forestal, Ministerio de Medio Ambiente, y Medio Rural y Marino, Madrid, Spain, 2009. View at: Google Scholar
  35. L. S. Glasgow and G. R. Matlack, “Prescribed burning and understory composition in a temperate deciduous forest, Ohio, USA,” Forest Ecology and Management, vol. 238, no. 1–3, pp. 54–64, 2007. View at: Publisher Site | Google Scholar
  36. J. Climent, R. Tapias, J. A. Pardos, and L. Gil, “Fire adaptations in the Canary Islands pine (Pinus canariensis),” Plant Ecology, vol. 171, no. 1-2, pp. 185–196, 2004. View at: Publisher Site | Google Scholar
  37. Gobierno de Canarias, Ley 12/1994, de 19 de diciembre, de Espacios Naturales de Canaria, Gobierno de Canarias, Boletín Oficial de Canarias no. 157, 1994.
  38. F. Grillo Delgado, D. Díaz Fababú, and J. Camaño Azcárate, III Curso Européo Avanzado de Manejo de Fuego, Las Palmas de Gran Canaria, 2009.
  39. A. L. Page, R. H. Miller, and D. R. Keeney, Methods of Soil Analysis: Part 2. Chemical and Microbiological Properties, ASA Monograph Number 9, ASA, Madison, Wis, USA, 1982.
  40. M. Anderson, R. Gorley, and K. Clarke, Permanova + for Primer: Guide to Software and Statistical Methods, Primer-E, Plymouth, UK, 2008.
  41. K. R. Clarke and R. M. Warwick, Change in Marine Communities: An Approach to Statistical Analysis and Interpretation, PRIMER-E, Plymouth, UK, 2nd edition, 2001.
  42. K. R. Clarke, “Non-parametric multivariate analyses of changes in community structure,” Australian Journal of Ecology, vol. 18, no. 1, pp. 117–143, 1993. View at: Publisher Site | Google Scholar
  43. J. Lepš and P. Šmilauer, Multivariate Analysis of Ecological Data Using CANOCO, Cambridge University Press, Cambridge, UK, 2003.
  44. C. J. F. Ter Braak and P. Šmilauer, CANOCO Reference Manual and User's Guide to Canoco for Windows: Software for Canonical Community Ordination (version 4), Microcomputer Power, Ithaca, NY, USA, 1998.
  45. M. J. Baeza and V. R. Vallejo, “Vegetation recovery after fuel management in Mediterranean shrublands,” Applied Vegetation Science, vol. 11, no. 2, pp. 151–158, 2008. View at: Publisher Site | Google Scholar
  46. J. C. Lake and M. R. Leishman, “Invasion success of exotic plants in natural ecosystems: the role of disturbance, plant attributes and freedom from herbivores,” Biological Conservation, vol. 117, no. 2, pp. 215–226, 2004. View at: Publisher Site | Google Scholar
  47. J. R. Arévalo and J. M. Fernández-Palacios, “Natural regeneration of Pinus canariensis Chr. Sm. Ex DC in Buch in forest plantations after thinning,” The Open Forest Science Journal, vol. 1, no. 1, pp. 54–60, 2008. View at: Publisher Site | Google Scholar
  48. K. L. Griffis, J. A. Crawford, M. R. Wagner, and W. H. Moir, “Understory response to management treatments in northern Arizona ponderosa pine forests,” Forest Ecology and Management, vol. 146, no. 1–3, pp. 239–245, 2001. View at: Publisher Site | Google Scholar
  49. C. García-Domínguez, Impacto del fuego en los procesos ecológicos relacionados con el mantenimiento de la diversidad en pinares repoblados de Pinus canariensis [Ph.D. thesis], University of La Laguna, La Laguna, Spain, 2010.
  50. J. R. Arévalo, J. M. Fernández-Palacios, M. J. Jiménez, and P. Gil, “The effect of fire intensity on the understorey species composition of two Pinus canariensis reforested stands in Tenerife (Canary islands),” Forest Ecology and Management, vol. 148, no. 1–3, pp. 21–29, 2001. View at: Publisher Site | Google Scholar
  51. C. García-Domínguez and J. M. Fernández-Palacios, “Effect of fire intensity on non-native plant species community in a Canarian pine forest three and eleven years after fire,” The Open Forest Science Journal, vol. 2, pp. 70–77, 2009. View at: Google Scholar
  52. L. Kokkoris, “Flowering and fruiting phenology of four herbaceous species of Leguminosae in aburned Pinus halepensis forest of Attica, Greece,” Journal of Mediterranean Ecology, vol. 1, pp. 193–200, 2000. View at: Google Scholar
  53. C. L. Wienk, C. H. Sieg, and G. R. McPherson, “Evaluating the role of cutting treatments, fire and soil seed banks in an experimental framework in ponderosa pine forests of the Black Hills, South Dakota,” Forest Ecology and Management, vol. 192, no. 2-3, pp. 375–393, 2004. View at: Publisher Site | Google Scholar
  54. J. R. Arévalo, A. Naranjo, J. M. Fernández-Palacios, and S. Fernández-Lugo, “Ecology and management of natural and reforested Canary Island pine stands,” in Woodlands: Ecology, Management and Conservation, Nova, 2011. View at: Google Scholar
  55. D. G. Simard, J. W. Fyles, D. Paré, and T. Nguyen, “Impacts of clearcut harvesting and wildfire on soil nutrient status in the Quebec boreal forest,” Canadian Journal of Soil Science, vol. 81, no. 2, pp. 229–237, 2001. View at: Publisher Site | Google Scholar
  56. D. J. Pennock and C. van Kessel, “Clear-cut forest harvest impacts on soil quality indicators in the mixedwood forest of Saskatchewan, Canada,” Geoderma, vol. 75, no. 1-2, pp. 13–32, 1997. View at: Publisher Site | Google Scholar
  57. G. Certini, “Effects of fire on properties of forest soils: a review,” Oecologia, vol. 143, no. 1, pp. 1–10, 2005. View at: Publisher Site | Google Scholar
  58. D. Gillon and M. Rapp, “Nutrient losses during a winter low-intensity prescribed fire in a Mediterranean forest,” Plant and Soil, vol. 120, no. 1, pp. 69–77, 1989. View at: Publisher Site | Google Scholar
  59. C. P. Giardina and C. C. Rhoades, “Clear cutting and burning affect nitrogen supply, phosphorus fractions and seedling growth in soils from a Wyoming lodgepole pine forest,” Forest Ecology and Management, vol. 140, no. 1, pp. 19–28, 2001. View at: Publisher Site | Google Scholar
  60. M. J. Gundale, T. H. DeLuca, C. E. Fiedler, P. W. Ramsey, M. G. Harrington, and J. E. Gannon, “Restoration treatments in a Montana ponderosa pine forest: effects on soil physical, chemical and biological properties,” Forest Ecology and Management, vol. 213, no. 1–3, pp. 25–38, 2005. View at: Publisher Site | Google Scholar
  61. M. C. Carter and C. D. Foster, “Prescribed burning and productivity in Southern Pine Forests: a review,” Forest Ecology and Management, vol. 191, no. 1–3, pp. 93–109, 2004. View at: Publisher Site | Google Scholar
  62. P. M. Fernandes and H. S. Botelho, “A review of prescribed burning effectiveness in fire hazard reduction,” International Journal of Wildland Fire, vol. 12, no. 2, pp. 117–128, 2003. View at: Publisher Site | Google Scholar
  63. G. Xanthopoulos, D. Caballero, M. Galante, D. Alexandrian, E. Rigolot, and R. Marzano, “Forest fuels management in Europe,” in USDA Forest Service Proceedings, pp. 29–46, 2006. View at: Google Scholar
  64. H. Hesseln, J. B. Loomis, and A. González-Cabán, “Comparing the economic effects of fire on hiking demand in Montana and Colorado,” Journal of Forest Economics, vol. 10, no. 1, pp. 21–35, 2004. View at: Publisher Site | Google Scholar
  65. R. T. Brown, J. K. Agee, and J. F. Franklin, “Forest restoration and fire: principles in the context of place,” Conservation Biology, vol. 18, no. 4, pp. 903–912, 2004. View at: Publisher Site | Google Scholar

Copyright © 2014 José Ramón Arévalo et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

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